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BACKGROUND: The present research aimed to develop and validate a standardised survey instrument for the assessment of patients' awareness of the quality of their oral hygiene performance. METHODS: A digital questionnaire was developed that assesses both patients' naïve self-perceptions of oral cleanliness (SPOCn) after tooth brushing and patients' perceptions after being informed how oral cleanliness may be captured in dentistry (SPOCd). Three studies (N = 56 adults, N = 66 adolescents and one of their parents, N = 24 university students) assessed the instrument's feasibility (patient reports), reliability (internal consistency), validity (correlation with other constructs; sensitivity to manipulation of actual tooth brushing), and the correlation with actual oral cleanliness after tooth brushing. RESULTS: All study groups accepted the questionnaire well; average answering times were less than 5 min. Cronbach's α exceeds 0.90; correlational analyses support the discriminant validity regarding oral hygiene related self-efficacy expectations and stages of change; manipulation of oral hygiene behaviour results in the expected changes of SPOC scores. Patients' SPOC correlate only moderately with actual oral cleanliness. The comparison between SPOCd scores and actual oral cleanliness indicate that they considerably overestimate their oral hygiene performance. CONCLUSIONS: The SPOC questionnaire is an easy-to-use, well-accepted, reliable and valid instrument for the assessment of patients' awareness of the quality of their oral hygiene for research and clinical purposes. The results of the questionnaire may help to reveal unrealistic self-perceptions of patients regarding their oral hygiene. It can raise their awareness of the need to improve their skills and/or efforts in this regard. Trial registration The third study was an interventional study and was registered in the appropriate national register ( www.drks.de ; ID: DRKS00018781; date of registration: 12/09/2019).
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Higiene Bucal , Escovação Dentária , Adulto , Adolescente , Humanos , Higiene Bucal/educação , Reprodutibilidade dos Testes , Escovação Dentária/métodos , Higiene , AutoeficáciaRESUMO
Computed-tomography-derived (CT-derived) polymesh surfaces are widely used in geometric morphometric studies. This approach is inevitably associated with decisions on scanning parameters, resolution, and segmentation strategies. Although the underlying processing steps have been shown to potentially contribute artefactual variance to three-dimensional landmark coordinates, their effects on measurement error have rarely been assessed systematically in CT-based geometric morphometric studies. The present study systematically assessed artefactual variance in landmark data introduced by the use of different voxel sizes, segmentation strategies, surface simplification degrees, and by inter- and intra-observer differences, and compared their magnitude to true biological variation. Multiple CT-derived surface variants of the anuran (Amphibia: Anura) pectoral girdle were generated by systematic changes in the factors that potentially influence the surface geometries. Twenty-four landmarks were repeatedly acquired by different observers. The contribution of all factors to the total variance in the landmark data was assessed using random-factor nested permanovas. Selected sets of Euclidean distances between landmark sets served further to compare the variance among factor levels. Landmark precision was assessed by landmark standard deviation and compared among observers and days. Results showed that all factors, except for voxel size, significantly contributed to measurement error in at least some of the analyses performed. In total, 6.75% of the variance in landmark data that mimicked a realistic biological study was caused by measurement error. In this landmark dataset, intra-observer error was the major source of artefactual variance followed by inter-observer error; the factor segmentation contributed < 1% and slight surface simplification had no significant effect. Inter-observer error clearly exceeded intra-observer error in a different landmark dataset acquired by six partly inexperienced observers. The results suggest that intra-observer error can potentially be reduced by including a training period prior to the actual landmark acquisition task and by acquiring landmarks in as few sessions as possible. Additionally, the application of moderate and careful surface simplification and, potentially, also the use of case-specific optimal combinations of automatic local thresholding algorithms and parameters for segmentation can help reduce intra-observer error. If landmark data are to be acquired by several observers, it is important to ensure that all observers are consistent in landmark identification. Despite the significant amount of artefactual variance, we have shown that landmark data acquired from microCT-derived surfaces are precise enough to study the shape of anuran pectoral girdles. Yet, a systematic assessment of measurement error is advisable for all geometric morphometric studies.
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Pontos de Referência Anatômicos/diagnóstico por imagem , Anuros/anatomia & histologia , Imageamento Tridimensional , Microtomografia por Raio-X , Animais , Esqueleto/diagnóstico por imagemRESUMO
The articulated appendages of arthropods are highly adaptable and potentially multifunctional, used for walking, swimming, feeding, prey capture, or other functions. Webspinners (Order Embioptera) are a paragon in this context. In contrast to other arthropods producing silk, they utilize their front feet for silk production. However, employing the same leg for alternative functions rather than for pure locomotion potentially imposes constraints and compromises. We here present morphological and experimental evidence for a "passive" pressure-induced silk spinning mechanism induced by external mechanical stimuli. Furthermore, we demonstrate that, as a consequence of the conflicting functions for their front feet, webspinners have evolved a unique style of walking that reduces the potentially problematic contact between silk ejectors and the substrate. Here we answer for the first time a long-term question within this enigmatic group of insects-how webspinners can use their front feet to spin their nanoscale silk. This knowledge may open the door for experimental studies on an artificial spinning process and for future utilization in applied fields of robotics or chemistry.
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Neópteros/fisiologia , Seda , Animais , Reação de Fuga , Feminino , Estimulação Física , Pressão , TatoRESUMO
In Odonata, a direct flight mechanism with specialized tendons evolved. One particular adaptation, the implementation of the rubber-like protein resilin in these cap tendons, might be of major importance. Although resilin was first described in one tendon of Odonata, to our knowledge no comprehensive study about the presence of resilin in the thorax exists yet. We investigated various species of Odonata, using µCT, dissection and fluorescence microscopy. Here we show a complete mapping of the odonatan pterothorax, regarding the presence of tendons and their properties. Thus, 20-21 cap tendons in the pterothorax of Odonata show the presence of resilin. While performing outstanding and often-aggressive flight manoeuvres, resilin can provide shock absorption against mechanical damage from strong impacts. It may further improve the wear and fatigue resistance owing to resilin's damping behaviour. Additionally, resilin in tendons can absorb and return kinetic energy to restore muscles to their original shape after contracting and help in maintaining self-oscillation of the flight muscles. Here, the material distribution within the direct flight system of Odonata and the biomechanical importance and possible function of resilin are discussed. These results are an important step towards the understanding of the complex form-material-function interplay of the insect cuticle.
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Odonatos , Animais , Voo Animal , Proteínas de Insetos , Insetos , Tendões , Asas de AnimaisRESUMO
The louse fly Crataerina pallida is an obligate blood-sucking ectoparasite of the common swift Apus apus As a result of reduction of the wings, C. pallida is unable to fly; thus, an effective and reliable attachment to their host's plumage is of utmost importance. The attachment system of C. pallida shows several modifications in comparison to that of other calyptrate flies, notably the large tridentate claws and the dichotomously shaped setae located on the pulvilli. Based on data from morphological analysis, confocal laser scanning microscopy, cryo-scanning electron microscopy and attachment force experiments performed on native (feathers) as well as artificial substrates (glass, epoxy resin and silicone rubber), we showed that the entire attachment system is highly adapted to the fly's lifestyle as an ectoparasite. The claws in particular are the main contributor to strong attachment to the host. Resulting attachment forces on feathers make it impossible to detach C. pallida without damage to the feathers or to the legs of the louse fly itself. Well-developed pulvilli are responsible for the attachment to smooth surfaces. Both dichotomously shaped setae and high setal density explain high attachment forces observed on smooth substrates. For the first time, we demonstrate a material gradient within the setae, with soft, resilin-dominated apical tips and stiff, more sclerotized bases in Diptera. The empodium seems not to be directly involved in the attachment process, but it might operate as a cleaning device and may be essential to maintain the functionality of the entire attachment system.
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Aves/parasitologia , Dípteros/anatomia & histologia , Plumas/parasitologia , Interações Hospedeiro-Parasita , Animais , Dípteros/fisiologia , Feminino , MasculinoRESUMO
BACKGROUND: The understanding of concerted movements and its underlying biomechanics is often complex and elusive. Functional principles and hypothetical functions of these complex movements can provide a solid basis for biomechanical experiments and modelling. Here a description of the cephalic anatomy of Pyrrhosoma nymphula (Zygoptera, Coenagrionidae) focusing on functional aspects of the mouthparts using micro computed tomography (µCT) is presented. RESULTS: We compared six different instars of the damselfly P. nymphula as well as one instar of the dragonfly Aeshna cyanea and Epiophlebia superstes each. In total 42 head muscles were described with only minor differences of the attachment points between the examined species and the absence of antennal muscle M. scapopedicellaris medialis (0an7) in Epiophlebia as a probable apomorphy of this group. Furthermore, the ontogenetic differences between the six larval instars are minor; the only considerable finding is the change of M. submentopraementalis (0la8), which is dichotomous in the early instars (I1,I2 and I3) with a second point of origin at the postero-lateral base of the submentum. This dichotomy is not present in any of the older instars studied (I6, middle-late and pen-ultimate). CONCLUSION: However, the main focus of the study herein, is to use these detailed morphological descriptions as basis for hypothetic functional models of the odonatan mouthparts. We present blueprint like description of the mouthparts and their musculature, highlighting the caused direction of motion for every single muscle. This data will help to elucidate the complex concerted movements of the mouthparts and will contribute to the understanding of its biomechanics not in Odonata only.
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The grey short-tailed opossum, Monodelphis domestica, has been an established research animal for more than five decades, but relatively, little is known about its thermophysiology. Here we studied core body temperature (T b) and metabolic rate (MR) of female adult M. domestica housed in the laboratory at an ambient temperature (T a) of 26 °C. In expanding previous reports, the average recorded core T b of M. domestica was 34.3 °C. The T b of an individual M. domestica can drop below 30 °C (minimal T b: 28.6 °C) accompanied by a reduction in MR of up to 52 % even while having ad libitum access to food. These findings demonstrate for the first time the presence of spontaneous torpor in M. domestica. Metabolic suppression at relatively high T a and T b furthermore broadens our perspective on the use of torpor as a metabolic strategy not just restricted to cold climates.
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Temperatura Alta , Monodelphis/fisiologia , Torpor/fisiologia , Animais , Temperatura Corporal , FemininoRESUMO
The arthropod cuticle offers strength, protection, and lightweight. Due to its limit in expandability, arthropods have to moult periodically to grow. While moulting is beneficial in terms of parasite or toxin control, growth and adaptation to environmental conditions, it costs energy and leaves the soft animal's body vulnerable to injuries and desiccation directly after ecdysis. To investigate the temporal change in sclerotization and pigmentation during and after ecdysis, we combined macrophotography, confocal laser scanning microscopy, scanning electron microscopy and histological sectioning. We analysed the tarsal and mandibular cuticle of the blue emperor dragonfly to compare the progress of tanning for structures that are functionally involved during emergence (tarsus/tarsal claws) with structures whose functionality is required much later (mandibles). Our results show that: (i) the tanning of the tarsal and mandibular cuticle increases during emergence; (ii) the tarsal cuticle tans faster than the mandibular cuticle; (iii) the mandibles tan faster on the aboral than on the oral side; and (iv) both the exo- and the endocuticle are tanned. The change in the cuticle composition of the tarsal and mandibular cuticle reflects the demand for higher mechanical stability of these body parts when holding on to the substrate during emergence and during first walking or hunting attempts.
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BACKGROUND: Among the winged insects (Pterygota) the Odonata (dragon- and damselflies) are special for several reasons. They are strictly aerial predators showing remarkable flight abilities and their thorax morphology differs significantly from that of other Pterygota in terms of the arrangement and number of muscles. Even within one individual the musculature is significantly different between the nymphal and adult stage. RESULTS: Here we present a comparative morphological investigation of the thoracic musculature of dragonfly (Anisoptera) nymphs. We investigated representatives of the Libellulidae, Aeshnidae and Cordulegasteridae and found 71 muscles: 19 muscles in the prothorax, 26 in the mesothorax and 27 in the metathorax. Nine of these muscles were previously unknown in Odonata, and for seven muscles no homologous muscles could be identified in the neopteran thorax. CONCLUSION: Our results support and extend the homology hypotheses for the thoracic musculatures of Odonata and Neoptera, thus supplementing our understanding of the evolution of Pterygota and providing additional characters for phylogenetic analyses comprising all subgroups of Pterygota.
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Evolução Biológica , Odonatos/anatomia & histologia , Odonatos/genética , Asas de Animais/anatomia & histologia , Animais , Voo Animal , Músculos/anatomia & histologia , Músculos/fisiologia , Odonatos/fisiologia , Filogenia , Asas de Animais/fisiologiaRESUMO
Prehensile raptorial forelegs are prey capturing and grasping devices, best known for praying mantises (Mantodea) within insects. They show strong morphological and behavioral adaptations toward a lifestyle as generalist arthropod predators. In the past, few species of Mantodea were investigated, concerning morphological variability of the raptorial forelegs. Especially the knowledge of foreleg anatomy in the light of functional and comparative morphology is scarce. Our comparative approach is based on the, for arthropods very common, "female-biased sexual size dimorphism" (SSD) that occurs in almost every Mantodea species. Within Mantodea, this SSD is likely leading to a shift of the exploited ecological niche between male and female individuals due to changes in, for example, the possible prey size; which might be reflected in the chosen ecomorphs. In this context, we analyzed the musculature of the raptorial forelegs of female and male specimens in five different species with varying SSD, using high-resolution microcomputed tomography and dissection. We were able to confirm the presence of 15 extrinsic and 15 intrinsic muscles-including one previously undescribed muscle present in all species. Thus, presenting a detailed description and illustrative three-dimensional anatomical visualization of the musculature in Mantodea. Interestingly, almost no observable differences were found, neither between species, nor between the sexes. Furthermore, we homologized all described muscles, due to their attachment points, to the comprehensive nomenclature established by Friedrich and Beutel (2008), discussed potential functionality of the muscles and possible homologies to the neuropteran Mantispa styriaca (Büsse et al., 2021) and the newly introduced leg nomenclature by Aibekova et al. (2022). By elucidating the anatomy, particularly in the context of functionality and SSD, our results complement previous knowledge of the raptorial forelegs, and facilitate a better understanding of the underlying biomechanical system of the predatory strike, and ultimately, a future comparison to other insect taxa.
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Mantódeos , Caracteres Sexuais , Animais , Masculino , Feminino , Mantódeos/fisiologia , Microtomografia por Raio-X , Insetos , Comportamento Predatório/fisiologiaRESUMO
Dragonfly and damselfly larvae (Insecta: Odonata) capture prey by rapid protraction of a raptorial mouthpart, based on a modified labium. Yet, in insects with biting-chewing mouthparts, the labium has an essential role in food handling. These two distinct functions -prey capturing and handling-lead to a mechanical problem in Odonata larvae: while the labium is always protracted in a straight line during prey capture, food handling requires more dexterity. In this study, we investigate the role of the labium in the feeding process and analyse the mechanics of the labial joints in the dragonfly larva Anax imperator. Our results show that the labium features a multiaxial joint connecting the basal segment (postmentum) and the head. During feeding, a combination of rotations around different axes is used to handle and orient prey, which is unique among biting-chewing mouthparts. Furthermore, we identified structures at the joint which likely restrict lateral motion during the predatory strike. Our results provide a further understanding of the unique prey-capturing apparatus of odonate larvae capable of controlling a 'switchable' multiaxial to a restricted monoaxial joint. This concept highlights the evolution of a highly modified raptorial mouthpart appendage where the degrees of freedom can be actively restricted to allow for the respectively needed functionality.
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Odonatos , Animais , Larva/fisiologia , Odonatos/fisiologia , Insetos , Mastigação , Comportamento Predatório/fisiologiaRESUMO
The species included in the genus Epiophlebia Calvert, 1903 represent an exception within Recent lineages - they do not belong to either dragonflies (Anisoptera) nor damselflies (Zygoptera). Nowadays, the genus is solely known from the Asian continent. Due to their stenoecious lifestyle, representatives of Epiophlebia are found in often very small relict populations in Nepal, Bhutan, India, Vietnam, China, North Korea, and Japan. We here present a taxonomic re-evaluation on the species status of Epiophlebiadiana Carle, 2012, known from the Sichuan province in China, supplemented with a morphological character mapping on a genetic tree to highlight synapomorphies of E.diana and E.laidlawi Tillyard, 1921. We conclude that E.diana is a junior synonym of E.laidlawi. Furthermore, we discuss the Recent distribution of the group, allowing for predictions of new habitats of representatives of this group.
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Barnacles rely heavily on their mobile cirri for food capture because of the sessile lifestyle. These filamentous food capturing devices are extended into the water current and perform undulating movements. Cuticular structures with corresponding musculature work together, to allow these highly repetitive movements. This paper studies the interplay between structure and function of the cirri using scanning electron microscopy (SEM), microcomputed tomography (µCT) and high-speed video recordings (HSV) in Amphibalanus improvisus. Barnacles use the external cuticle structures (denticles and setae) for efficient grooming, food capturing, and providing support for the muscular system responsible for movement control. Entanglement of the cirri during extension is probably avoided through an interlocking of the serrate setae on cirri IV-VI, creating a "zipper-like" effect, which was recorded here using the HSV. We analyzed the muscular arrangement using µCT and found a new flexor muscle in both the endo- and exopod of cirrus II. Supported by the intrinsic cirral muscles, the new flexor muscles may provide variable movements of the anterior cirri (cirri I-III), which is important for further food handling. Our results provide a foundation for further comparative studies of the feeding apparatus of barnacles and for possible implications in the area of bio-inspired robotics.
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Thoracica , Animais , Thoracica/anatomia & histologia , Thoracica/fisiologia , Microtomografia por Raio-X , Microscopia Eletrônica de Varredura , Músculos , ÁguaRESUMO
Insects evolved differently specialized mouthparts. We study the mouthparts of adult Anax imperator, one of the largest odonates found in Central Europe. Like all adult dragonflies, A. imperator possesses carnivorous-type of biting-chewing mouthparts. To gain insights into the feeding process, behavior and kinematics, living specimens were filmed during feeding using synchronized high-speed videography. Additionally, the maximum angles of movement were measured using a measuring microscope and combined with data from micro-computed tomography. The resulting visualizations of the 3D-geometry of each mouthpart were used to study their anatomy and complement the existing descriptive knowledge of muscles in A. imperator to date. Furthermore, confocal laser scanning microscopy-projections allow for estimation of differences in the material composition of the mouthparts' cuticle. By combining all methods, we analyze possible functions and underlying biomechanics of each mouthpart. We also analyzed the concerted movements of the mouthparts; unique behavior of the mouthparts during feeding is active participation by the labrum and distinct movement by the maxillary laciniae. We aim to elucidate the complex movements of the mouthparts and their functioning by combining detailed information on (1) in vivo movement behavior (supplemented with physiological angle approximations), (2) movement ability provided by morphology (morphological movement angles), (3) 3D-anatomy, and (4) cuticle composition estimates.
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Odonatos , Animais , Fenômenos Biomecânicos , Insetos , Mastigação , Microtomografia por Raio-XRESUMO
Hippoboscidae and Nycteribiidae of the dipteran superfamily Hippoboscoidea are obligate ectoparasites, which feed on the blood of different mammals. Due to their limited flight capability, the attachment system on all tarsi is of great importance for a secure grasp onto their host and thus for their survival. In this study, the functional morphology of the attachment system of two hippoboscid species and two nycteribiid species was compared in their specificity to the host substrate. Based on data from scanning electron microscopy and confocal laser scanning microscopy, it was shown that the attachment systems of both Hippoboscidae and Nycteribiidae (Hippoboscoidea) differ greatly from that of other calyptrate flies and are uniform within the respective families. All studied species have an attachment system with two monodentate claws and two pulvilli. The claws and pulvilli of the Hippoboscidae are asymmetric, which is an adaptation to the fur of even-toed ungulates (Artiodactyla). The fur of these mammals possesses both, thinner woolen and thicker coat hair; thus, the asymmetry of the attachment system of the hippoboscid species enables a secure attachment to all surfaces of their hosts. The claws and pulvilli of the nyceribiid species do not show an asymmetry, since the fur of their bat (Chiroptera) hosts consists of hairs with the same thickness. The claws are important for the attachment to mammals' fur, because they enable a secure grip by mechanical interlocking of the hairs through the claws. Additionally, well-developed pulvilli are able to attach on thicker hairs of Artiodactyla or on smooth substrates such as the skin.
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Quirópteros , Dípteros , Ftirápteros , Animais , CabeloRESUMO
The biomechanics underlying the predatory strike of dragonfly larvae is not yet understood. Dragonfly larvae are aquatic ambush predators, capturing their prey with a strongly modified extensible mouthpart. The current theory of hydraulic pressure being the driving force of the predatory strike can be refuted by our manipulation experiments and reinterpretation of former studies. Here, we report evidence for an independently loaded synchronized dual-catapult system. To power the ballistic movement of a single specialized mouthpart, two independently loaded springs simultaneously release and actuate two separate joints in a kinematic chain. Energy for the movement is stored by straining an elastic structure at each joint and, possibly, the surrounding cuticle, which is preloaded by muscle contraction. As a proof of concept, we developed a bioinspired robotic model resembling the morphology and functional principle of the extensible mouthpart. Understanding the biomechanics of the independently loaded synchronized dual-catapult system found in dragonfly larvae can be used to control the extension direction and, thereby, thrust vector of a power-modulated robotic system.
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Odonatos/fisiologia , Robótica/instrumentação , Animais , Fenômenos Biomecânicos , Materiais Biomiméticos , Comportamento Alimentar/fisiologia , Larva/anatomia & histologia , Larva/fisiologia , Modelos Biológicos , Boca/anatomia & histologia , Boca/fisiologia , Movimento/fisiologia , Odonatos/anatomia & histologia , Comportamento Predatório/fisiologia , Robótica/estatística & dados numéricosRESUMO
Sandy pitfall traps of antlions are elaborate constructions to capture prey. Antlions exploit the interactions between the particles in their habitat and build a stable trap. This trap is close to the unstable state; prey items will slide towards the centre-where the antlion ambushes-when entering the trap. This is efficient but requires permanent maintenance. According to the present knowledge, antlions throw sand, mainly to cause sandslides towards the centre of the pit. We hypothesized that: (i) sand-throwing causes sandslides towards the centre of the pit and (ii) sand-throwing constantly maintains the pitfall trap and thus keeps its efficiency high. Using laboratory experiments, as well as finite-element analysis, we tested these hypotheses. We show, experimentally and numerically, that sand that accumulates at the centre of the pit will be removed continuously by sand-throwing, this maintenance is leading to slope condition close to an unstable state. This keeps the slope angle steep and the efficiency of the trap constant. Furthermore, the resulting sandslides can relocate the trapped prey towards the centre of the pit. This study adds further insights from specific mechanical properties of a granular medium into the behavioural context of hunting antlion larvae.
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Insetos , Areia , Animais , Ecossistema , Larva , Comportamento PredatórioRESUMO
Analysing the motion of animals, especially at high speeds, is often challenging. Motion tracking software needs to deal with a variety of visual contexts, variable lighting conditions, heterogeneous backgrounds and even background movements. Here we present motion tracking via the easy to use and constantly updated Adobe After Effects software - which is often included in software packages most researchers are already using. The provided custom-made Javascript allows for easy exporting of tracking coordinates. Furthermore, some examples for analysing the obtained data in the open source statistical software 'R' will provide reference points, even for an unexperienced user. We present a step-by-step guide of the methodology using high-speed video recordings of locust jumps and additionally validate this method by successful tracking of simulated data under defined subpar filming conditions. This simulated data allows experienced users to compare the tracking software in use with the here presented workflow to weigh the advantages and disadvantages of any motion tracking software on the market.
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Gafanhotos/fisiologia , Atividade Motora/fisiologia , Gravação em Vídeo/métodos , Animais , Comportamento Animal , Fenômenos Biomecânicos , Processamento de Imagem Assistida por Computador , SoftwareRESUMO
Presenting your research in the proper light can be exceptionally challenging. Meanwhile, dome illumination systems became a standard for micro- and macrophotography in taxonomy, morphology, systematics and especially important in natural history collections. However, proper illumination systems are either expensive and/or laborious to use. Nowadays, 3D-printing technology revolutionizes lab-life and will soon find its way into most people's everyday life. Consequently, fused deposition modelling printers become more and more available, with online services offering personalized printing options. Here, we present a 3D-printed, scalable, low-cost and modular LED illumination dome system for scientific micro- and macrophotography. We provide stereolithography ('.stl') files and print settings, as well as a complete list of necessary components required for the construction of three differently sized domes. Additionally, we included an optional iris diaphragm and a sliding table, to arrange the object of desire inside the dome. The dome can be easily scaled and modified by adding customized parts, allowing you to always present your research object in the best light.
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Moulting, especially in 'hemimetabolous' insects that emerge upside down, is a crucial moment in their live. Losing their attachment during this situation can be fatal for survival. We here studied the emergence of dragonfly adults, describe structures involved in larval attachment to the substrate, and biomechanically test the pull-off forces of exuviae to natural substrates. Confocal laser scanning microscopy and scanning electron microscopy were used to describe both morphology and material composition of the leg cuticle of Anax imperator larvae. The results show that the combination of morphological and behavioral adaptations provides reliable anchorage of exuviae to the substrates. We determined a safety factor of 14, and demonstrated that this staggered safety system experiencing several unlocking and relocking events withstand multiple disturbances before the entire exuvia is completely detaches. This furthers our understanding of interlocking and anchorage of insects in general and may allow for future applications.