Lepidoptera demonstrate the relevance of Murray's Law to circulatory systems with tidal flow.

BACKGROUND: Murray's Law, which describes the branching architecture of bifurcating tubes, predicts the morphology of vessels in many amniotes and plants. Here, we use insects to explore the universality of Murray's Law and to evaluate its predictive power for the wing venation of Lepidoptera, one of the most diverse insect orders. Lepidoptera are particularly relevant to the universality of Murray's Law because their wing veins have tidal, or oscillatory, flow of air and hemolymph. We examined over one thousand wings representing 667 species of Lepidoptera. RESULTS: We found that veins with a diameter above approximately 50 microns conform to Murray's Law, with veins below 50 microns in diameter becoming less and less likely to conform to Murray's Law as they narrow. The minute veins that are most likely to deviate from Murray's Law are also the most likely to have atrophied, which prevents efficient fluid transport regardless of branching architecture. However, the veins of many taxa continue to branch distally to the areas where they atrophied, and these too conform to Murray's Law at larger diameters (e.g., Sesiidae). CONCLUSIONS: This finding suggests that conformity to Murray's Law in larger taxa may reflect requirements for structural support as much as fluid transport, or may indicate that selective pressures for fluid transport are stronger during the pupal stage-during wing development prior to vein atrophy-than the adult stage. Our results increase the taxonomic scope of Murray's Law and provide greater clarity about the relevance of body size.

X-ray computed tomography study of the flight-adapted tracheal system in the blowfly Calliphora vicina, analysing the ventilation mechanism and flow-directing valves.

Following the discovery of flight motor-driven unidirectional gas exchange with rising PO2  in the blowfly, X-ray computed tomography (CT) was used to visualize the organization of the tracheal system in the anterior body with emphasis on the arrangement of the pathways for airflow. The fly's head is preferentially supplied by cephalic tracheae originating from the ventral orifice of the mesothoracic spiracle (Sp1). The respiratory airflow during flight is a by-product of cyclic deformations of the thoracic box by the flight muscles. The air sacs below the tergal integument (scutum and scutellum) facilitate the respiratory airflow: the shortening of the thorax turns the scutellum and the wings downward and the scutum upward with a volume increase in the scutal air sacs. The resulting negative pressure sucks air from Sp1 through special tracheae towards the scutal air sacs. The airflow is directed by two valves that open alternately: (1) the hinged filter flaps of the metathoracic spiracles (Sp2) are passively pushed open during the upstroke by the increased tracheal pressure, thereby enabling expiration; (2) a newly described tracheal valve-like septum behind the regular spiracular valve lids of Sp1 opens passively and air is sucked in through Sp1 during the downstroke and prevents expiration by closing during the upstroke. This stabilizes the unidirectional airflow. The tracheal volume of the head, thorax and abdomen and their mass were determined. Despite the different anatomy of birds and flies, the unidirectional airflow reveals a comparable efficiency of the temporal throughput in flies and hummingbirds.

Structure of the thoracic spiracular valves and their contribution to unidirectional gas exchange in flying blowflies Calliphora vicina.

The operation of the thoracic spiracular valves was analysed using anatomical and physiological techniques. Dense spiracular filter trichomes impede a diffusive gas exchange. However, the hinged posterior filter flap of the metathoracic spiracle (Sp2) opens passively during upstroke of the wings and closes by the suction of the sub-atmospheric tracheal pressure during the downstroke, which supports a unidirectional respiratory airflow. The action of the interior spiracular valve lids was recorded by photocell sensors oriented above the enlarged spiracles and projected onto the screen of a video camera. The thoracic spiracles opened much quicker (approximately 0.1 s) than they closed (1 s), suggesting that the spiracular muscles are openers, as confirmed by experimental induction of muscle contraction. Simultaneous photocell measurement revealed that the first and second thoracic spiracles act concordantly. At rest, the spiracles were mostly closed or only slightly open (<1%). During intermittent short flights, the valves opened wide at the start of the flight for a short time, and in many cases opened again after the flight ended. Often, the opening was wider after the flight ended than during the flight itself. During long spontaneous continuous flight phases (up to 2 h), the valves were only slightly open (<5%), widening shortly after transient increases of wing stroke intensity. It is an amazing paradox that the spiracles were only slightly open when the requirement for O2 was high during sustained flight. The advantage of generating sub-atmospheric pressure, supporting a unidirectional airflow with a PO2  increase above the resting level, is discussed.

Flight-motor-driven respiratory airflow increases tracheal oxygen to nearly atmospheric level in blowflies (Calliphora vicina).

It is widely accepted that an efficient oxygen supply and removal of CO2 in small flying insects are sufficiently performed by diffusion with open spiracles. This paper shows that in the tethered flying blowfly, gas exchange occurs by autoventilation and unidirectional airflow. The air is inspired through the mesothoracic spiracles (Sp1) during the downstroke of the wings and is expired through the metathoracic spiracles (Sp2) during the upstroke. This directed airflow through the thoracic tracheal system was documented by pre-atrial pressure measurements at the Sp1 and Sp2, revealing a sub-atmospheric mean pressure at the Sp1 and an over-atmospheric mean pressure at the Sp2. In the mesothoracic air sacs, the mean pressure is sub-atmospheric, conditioned by the only slightly open spiracles. In a split flow-through chamber experiment, the CO2 released through the Sp2 confirmed this unidirectional respiratory gas flow, implicating an inner tracheal valve. In the thoracic tracheal system, the PO2  during flight exceeds the high resting PO2  by 1-2 kPa, reaching nearly atmospheric values. In the abdominal large air sacs, the PO2  drops during flight, probably due to the accumulation of CO2. Periodic heartbeat reversals continue during flight, with a higher period frequency than at rest, supporting the transport of CO2 via the haemolymph towards the metathoracic tracheae and abdominal air sacs.

Periodic heartbeat reversals cause cardiogenic inspiration and expiration with coupled spiracle leakage in resting blowflies, Calliphora vicina.

Respiration in insects is thought to be independent of the circulatory system because insects typically lack respiratory pigments and because oxygen transport occurs in the gaseous phase through a ramified tracheal system by diffusion and convection directly to the tissues. In the blowfly, as in other insects with periodic heartbeat reversal, the haemolymph is periodically shifted between the anterior body and abdomen, exerting alternating pressure changes on the compliant tracheae in the thorax and in the abdomen. Simultaneous pressure and O2 optode measurements show that, during negative pressure periods, the tracheal partial pressure of oxygen (PO2) increases by 0.5 kPa. In the quiescent fly, tracheal PO2 is rather high (17.5-18.9 kPa), although the thoracic spiracles remain constricted. Microscopic video recordings and reflectance measurements revealed that the dorsal soft edges of the valve lips of the second spiracle leave a very small leak, which is passively widened during backward pulses of the heart. Thus, negative pressure, combined with increased leakage of the spiracle Sp2 valve enable inspiration in the thorax. The positive pressure periods are correlated with a new type of convective CO2 micro-bursts as shown in flow-through measurements. The bulk of the CO2 is, however, released after longer interbursts in macro-bursts with actively opening valves reminiscent of the open phase in a cyclic gas exchange. When the valves open, the PO2 in the thoracic air sacs unexpectedly drops by a mean of 2.75±1.09 kPa, suggesting a displacement of O2 by the transient accumulation of CO2 in the tracheal system before its release.

Hypoxia-induced compression in the tracheal system of the tobacco hornworm caterpillar, Manduca sexta.

Abdominal pumping in caterpillars has only been documented during molting. Using synchrotron X-ray imaging in conjunction with high-speed flow-through respirometry, we show that Manduca sexta caterpillars cyclically contract their bodies in response to hypoxia, resulting in significant compressions of the tracheal system. Compression of tracheae induced by abdominal pumping drives external gas exchange, as evidenced by the high correlation between CO2 emission peaks and body movements. During abdominal pumping, both the compression frequency and fractional change in diameter of tracheae increased with body mass. However, abdominal pumping and tracheal compression were only observed in larger, older caterpillars (>0.2 g body mass), suggesting that this hypoxic response increases during ontogeny. The diameters of major tracheae in the thorax increased isometrically with body mass. However, tracheae in the head did not scale with mass, suggesting that there is a large safety margin for oxygen delivery in the head in the youngest animals. Together, these results highlight the need for more studies of tracheal system scaling and suggest that patterns of tracheal investment vary regionally in the body.

Coordinated ventilation and spiracle activity produce unidirectional airflow in the hissing cockroach, Gromphadorhina portentosa.

Insects exchange respiratory gases via an extensive network of tracheal vessels that open to the surface of the body through spiracular valves. Although gas exchange is known to increase with the opening of these spiracles, it is not clear how this event relates to gas flow through the tracheal system. We examined the relationship between respiratory airflow and spiracle activity in a ventilating insect, the hissing cockroach, Gromphadorhina portentosa, to better understand the complexity of insect respiratory function. Using simultaneous video recordings of multiple spiracular valves, we found that abdominal spiracles open and close in unison during periods of ventilation. Additionally, independent recordings of CO2 release from the abdominal and thoracic regions and observations of hyperoxic tracer gas movement indicate that air is drawn into the thoracic spiracles and expelled from the abdominal spiracles. Our video recordings suggest that this unidirectional flow is driven by abdominal contractions that occur when the abdominal spiracles open. The spiracles then close as the abdomen relaxes and fills with air from the thorax. Therefore, the respiratory system of the hissing cockroach functions as a unidirectional pump through the coordinated action of the spiracles and abdominal musculature. This mechanism may be employed by a broad diversity of large insects that respire by active ventilation.

Mass and volume growth of an insect tracheal system within a single instar.

Organisms must accommodate oxygen delivery to developing tissues as body mass increases during growth. In insects, the growth of the respiratory system has been assumed to occur only during molts, whereas body mass and volume increase during the larval stages between molts. This decouples whole-body growth from the growth of the oxygen supply system. This assumption is derived from the observation that the insect respiratory system is an invagination of the exoskeleton, which must be shed during molts for continued growth to occur. Here, we provide evidence that this assumption is incorrect. We found that the respiratory system increases substantially in both mass and volume within the last larval instar of Manduca sexta larvae, and that the growth of the respiratory system changes with diet quality, potentially as a consequence of shifting metabolic demands.

Ideal endotracheal intubation depth at the vocal-cord level to avoid single-lung intubation using the percentage ratio of the tracheal length to body height.

INTRODUCTION: Our previous study revealed racial differences in the tracheal length of cardiac paediatric patients between Germany and Japan. The current study was conducted in two stages, aiming to determine whether the tracheal length differs between cardiac and non-cardiac paediatric patients and whether the results could also be generalised to adults. MATERIAL AND METHODS: The first stage was a retrospective observational evaluation of 335 cardiac and 275 non-cardiac paediatric patients in Japan. The tracheal length, the distance between the vocal cords and carina tracheae, was measured on preoperative chest radiographs taken in the supine position. The second stage was a validation process including 308 Japanese patients. Endotracheal intubation was performed based on the results of the first-stage investigation. RESULTS: It was revealed that the tracheal length ranged from 7 to 11% of the body height in both the cardiac and non-cardiac Japanese paediatric patients. None of 308 Japanese paediatric and adult patients underwent single-lung intubation after the endotracheal tube was inserted at a depth of 7% of the body height at the vocal-cord level, corresponding to the minimum tracheal length for Japanese patients. The distance between the endotracheal tube tip and carina tracheae on postoperative chest radiographs was generally less than 4% of the body height across all paediatric and adult Japanese patients. CONCLUSIONS: The current study demonstrated that endotracheal intubation avoiding single-lung intubation can be achieved by inserting endotracheal tubes to the minimum tracheal length for a specific ethnic group at the vocal-cord level in paediatric patients, including neonates and premature infants, as well as adults.

Ideal Depth of Endotracheal Intubation at the Vocal Cord Level in Pediatric Patients Considering Racial Differences in Tracheal Length.

Numerous formulas that can predict endotracheal intubation depth at the corner of the mouth or the nasal wing of patients have been reported, even though the oral and nasal cavity anatomies differ among patients. Therefore, the purpose of this study was to derive a simple and reliable formula to predict the ideal endotracheal tube insertion depth at the vocal cord level in pediatric patients. The current study was conducted as a retrospective observational study, involving 425 and 335 cardiac pediatric patients in Germany and Japan, respectively, and aimed to determine a formula for predicting tracheal length and ideal depth of endotracheal intubation at the vocal cord level in pediatric patients. The distance between the vocal cords and the carina tracheae was defined as the tracheal length, and was measured on preoperative chest radiographs obtained in the supine position. The tracheal length in cardiac pediatric patients ranged from 6 to 10% of the body height in Germany and from 7 to 11% in Japan. This study revealed racial differences in the tracheal length, that is, in the ideal depth of endotracheal intubation at the vocal cord level. This study suggests that an adequate endotracheal intubation depth can be achieved by inserting endotracheal tubes at the vocal cord level with the minimum tracheal length of each racial group in pediatric patients, for example, 6% and 7% of the body height in Europeans and Asians, respectively. If the endotracheal tube inserted with this method appears to be shallow on chest radiographs, this does not represent an increased risk of accidental extubation, due to an excessively short intubation depth, because the minimum tracheal length for each racial group is considered. That is, it is not due to the endotracheal tube insertion length, but is likely due to the tracheal length of the patient, who has a relatively long tracheal length in the racial group.

A review of the hexapod tracheal system with a focus on the apterygote groups.

Respiratory systems are key innovations for the radiation of terrestrial arthropods. It is therefore surprising that there is still a considerable lack of knowledge. In this review of the available information on tracheal systems of hexapods (with a focus on the apterygote lineages Protura, Collembola, Diplura, Archaeognatha and Zygentoma), we summarize available data on the spiracles (number, position and morphology), the shape and variability of tracheal branching patterns including anastomoses, the tracheal fine structure and the respiratory proteins. The available data are strongly fragmented, and information for most subgroups is missing. In various cases, individual observations for one species account for the knowledge of the entire order. The available data show that there are strong differences between but also within apterygote orders. We conclude that the available data are insufficient to derive detailed conclusions on the hexapod ground plan and outline the possible evolutionary scenarios for the tracheal system in this group.

Evidence for wing development in the Late Palaeozoic Palaeodictyoptera revisited.

The numerous fossil specimens described as consecutive series of different larval stages of two species, Tchirkovaea guttata and Paimbia fenestrata (Palaeodictyoptera: Tchirkovaeidae), were reinvestigated with emphasis on comparing the development and growth of their wings with that of the wings of a recent mayfly, Cloeon dipterum. This unique fossil material was for a long time considered as undisputed evidence for an unusual type of wing development in Palaeozoic insects. The original idea was that the larvae of Palaeodictyopterida had wings, which were articulated and fully movable in their early stages of postembryonic development and that these gradually enlarging wings changed their position from longitudinal to perpendicular to the body axis. Moreover, the development of wings was supposed to include two or more subimaginal instars, implying that the fully winged instars moulted several times during their postembryonic development. The results of the present study revealed that there is no evidence that this series of nymphal, subimaginal and imaginal wings provide support for the original idea of wing development in Palaeozoic insects. On the contrary, our results indicate, that the supposed palaeodictyopteran larval wings are in fact wing pads with a wing developing inside the cuticular sheath as in recent hemimetabolous insects. Moreover, this study newly reinterpreted the wing pad base of Parathesoneura carpenteri and confirmed the presence of nygma like structures on wings and wing pads of palaeodictyopteran Tchirkovaeidae.

Proteomics reveals localization of cuticular proteins in Anopheles gambiae.

Anopheles gambiae devotes over 2% of its protein coding genes to its 298 structural cuticular proteins (CPs). This paper provides new LC-MS/MS data on two adult structures, proboscises and palps, as well as three larval samples - 4th instar larvae, just their terminal segment, and a preparation enriched in their tracheae. These data were combined with our previously published results of proteins from five other adult structures, whole adults, and two preparations chosen for their relatively clean cuticle, the larval head capsules left behind after ecdysis and the pupal cuticles left behind after adult eclosion. Peptides from 28 CPs were recovered in all adult structures; 24 CPs were identified for the first time, 6 of these were members of the TWDL family. Most newly identified proteins came from the larval sources. Based solely on peptide recovery, from our data and from other investigators, most available on VectorBase, there were only 4 CPs that were restricted to a single adult structure. More were restricted to a single metamorphic stage, 14 in larvae, 0 in pupae and 32 in adults. Expression data from our earlier RT-qPCR studies reduces these numbers. Charting restriction of CPs to stage or structure is a step forward in establishing their specific roles.

Viviparity in the dermapteran Arixenia esau: respiration inside mother's body requires both maternal and larval contribution.

Earwigs (Dermaptera) use different strategies to increase their reproductive success. Most species lay eggs; however, viviparity of the matrotrophic type has been reported in two groups: Hemimeridae and Arixeniidae. In Arixeniidae, offspring develop in two separate places: inside an ovary (the intraovarian phase) and within a uterus (the intrauterine phase). Both morphological and physiological aspects of viviparity in Arixeniidae have begun to be unraveled only recently. Here, we characterize how the first instar larvae of Arixenia esau, developing inside the mother's reproductive system, manage respiration and gas exchange. Using modern light and electron microscopy techniques as well as immunological approach, we provide a detailed account of the maternal and larval tissue interactions during the intrauterine development. We demonstrate that respiration in the Arixenia first instar larvae relies on the extensive tracheal system of the mother as well as a respiratory pigment (hemocyanin) present within the body cavity of the larvae. Our results indicate that the larval fat body tissue is the likely place of the hemocyanin synthesis. Our study shows that characteristic cone-shaped lobes of the outgrowths located on the larval abdomen are a part of a placenta-like organ and mediate the gas exchange between the maternal and larval organisms. Based on the obtained results, we propose that Arixenia esau evolved a unique biphasic system supporting respiration of the first instar larvae during their development inside the mother's reproductive tract.

A new species of the armored spider genus Caraimatta Lehtinen, 1981 from Colombia (Araneae: Synspermiata: Tetrablemmidae).

A new species of the armored spider genus Caraimatta Lehtinen, 1981 from Colombian Tropical dry forest fragments is described and illustrated: Caraimatta brescoviti sp. nov. (based on male and female) from Bolivar and Sucre departments, representing the first record of the genus from Colombia. Additionally, photographs of Monoblemma muchmorei Shear, 1978 (other tetrablemmid species previously recorded from the country) are given. A map with the known distribution and an identification key for males and females of the Caraimatta species are also included.

Developmental plasticity and stability in the tracheal networks supplying Drosophila flight muscle in response to rearing oxygen level.

While it is clear that the insect tracheal system can respond in a compensatory manner to both hypoxia and hyperoxia, there is substantial variation in how different parts of the system respond. However, the response of tracheal structures, from the tracheoles to the largest tracheal trunks, have not been studied within one species. In this study, we examined the effect of larval/pupal rearing in hypoxia, normoxia, and hyperoxia (10, 21 or 40kPa oxygen) on body size and the tracheal supply to the flight muscles of Drosophila melanogaster, using synchrotron radiation micro-computed tomography (SR-µCT) to assess flight muscle volumes and the major tracheal trunks, and confocal microscopy to assess the tracheoles. Hypoxic rearing decreased thorax length whereas hyperoxic-rearing decreased flight muscle volumes, suggestive of negative effects of both extremes. Tomography at the broad organismal scale revealed no evidence for enlargement of the major tracheae in response to lower rearing oxygen levels, although tracheal size scaled with muscle volume. However, using confocal imaging, we found a strong inverse relationship between tracheole density within the flight muscles and rearing oxygen level, and shorter tracheolar branch lengths in hypoxic-reared animals. Although prior studies of larger tracheae in other insects indicate that axial diffusing capacity should be constant with sequential generations of branching, this pattern was not found in the fine tracheolar networks, perhaps due to the increasing importance of radial diffusion in this regime. Overall, D. melanogaster responded to rearing oxygen level with compensatory morphological changes in the small tracheae and tracheoles, but retained stability in most of the other structural components of the tracheal supply to the flight muscles.

Morphological responses to feeding in ticks (Ixodes ricinus).

BACKGROUND: Ticks can survive long periods without feeding but, when feeding, ingest large quantities of blood, resulting in a more than 100-fold increase of body volume. We study morphological adaptations to changes in opisthosoma volume during feeding in the castor bean tick, Ixodes ricinus. We aim to understand the functional morphological features that accommodate enormous changes in volume changes. METHODS: Using light and electron microscopy, we compare the cuticle and epidermis of the alloscutum, the epithelium of the midgut diverticula, and the tracheae of adult female ticks when fasting, semi-engorged, and fully engorged. RESULTS: Our results add to an existing body of knowledge that the area of the epidermis increases by cellular differentiation, cellular hypertrophy, and changes in the shape of epithelial cells from pseudostratified to single layered prismatic in semi-engorged ticks, and to thin squamous epithelium in fully engorged ticks. We did not find evidence for cell proliferation. The midgut diverticula accommodate the volume increase by cellular hypertrophy and changes in cell shape. In fully engorged ticks, the epithelial cells of the midgut diverticula are stretched to an extremely thin, squamous epithelium. Changes in size and shape (and cell divisions) contribute to the accommodation of volume changes. Tracheae do not increase in size, but extend in length, thus following the volume changes of the opisthosoma in feeding ticks to secure oxygen supply to the internal organs. CONCLUSIONS: Changes of epithelial tissue configuration in the epidermis and the midgut diverticula are described as important components of the morphological response to feeding in ticks. We provide evidence for a previously unknown mechanism hosted in the endocuticle of the tracheae that allows the tracheae of castor bean ticks to expand when the body volume increases and the distance between the respiratory spiracle and the oxygen demanding tissue enlarges. This is the first report of expandable tracheae in arthropods.

Divergent roles of the Drosophila melanogaster globins.

In contrast to long-held assumptions, the gene repertoire of most insects includes hemoglobins. Analyses of the genome of the fruitfly Drosophila melanogaster identified three distinct hemoglobin genes (glob1, glob2, and glob3). While glob1 is predominantly associated with the tracheal system and fat body, glob2 and glob3 are almost exclusively expressed in the testis. The physiological role of globins in Drosophila is uncertain. Here, we studied the functions of the three globins in a cell culture system. Drosophila Schneider 2 (S2) cells were stably transfected with each of the three globins and the empty vector as control. Under hypoxia (1% atmospheric O2), only glob1 overexpression enhanced the activity of mitochondrial oxidases and the ATP content. However, the positive effect of glob1 expression disappeared after 24h hypoxia, suggesting metabolic adaptations of the S2 cells. glob2 and glob3 had no positive effect on hypoxia-survival. After application of oxidative stress by H2O2, glob2 dramatically enhanced the viability of S2 cells. Evaluation of the intracellular localization of the globins using specific antibodies and green fluorescent protein-fusion constructs suggested that glob1 and glob2 most likely reside in the cytoplasm, while glob3 is associated with structures that may represent parts of the intracellular transport machinery. In silico analyses of public RNA-Seq data from different developmental stages provided that glob1 is co-expressed with genes of the aerobic energy metabolism, while glob2 and glob3 expression can be related to spermatogenesis and reproduction. Together, the results indicate divergent functions of the Drosophila globins: glob1 may play a role in the O2-dependent metabolism while glob2 may protect spermatogenesis from reactive oxygen species.

Supply and demand: How does variation in atmospheric oxygen during development affect insect tracheal and mitochondrial networks?

Atmospheric oxygen is one of the most important atmospheric component for all terrestrial organisms. Variation in atmospheric oxygen has wide ranging effects on animal physiology, development, and evolution. This variation in oxygen has the potential to affect both respiratory systems (the supply side) and mitochondrial networks (the demand side) in animals. Insect respiratory systems supplying oxygen to tissues in the gas phase through blind ended tracheal systems are particularly susceptible to this variation. While the large conducting tracheae have previously been shown to respond developmentally to changes in rearing oxygen, the effect of oxygen on the tracheolar network has been relatively unexplored, especially in adult insects. Similarly, mitochondrial networks that meet energy demand in insects and other animals are dynamic and their enzyme activities have been shown to vary in the presence of oxygen. These two systems together should be under selective pressure to meet the aerobic metabolic requirements of insects. To test this hypothesis, we reared Mito-YFP Drosophila under three different oxygen concentrations hypoxia (12%), normoxia (21%), and hyperoxia (31%) and imaged their tracheolar and mitochondrial networks within their flight muscle using confocal microscopy. In terms of oxygen supply, hypoxia increased mean (mid-length) tracheolar diameters, tracheolar tip diameters, the number of tracheoles per main branch and affected tracheal branching patterns, while the opposite was observed in hyperoxia. In terms of oxygen demand, hypoxia increased mitochondrial investment and mitochondrial to tracheolar volume ratios; while the opposite was observed in hyperoxia. Generally, hypoxia had a stronger effect on both systems than hyperoxia. These results show that insects are capable of developmentally changing investment in both their supply and demand networks to increase overall fitness.

The forewing of the Aphis fabae (Scopoli 1763) (Hemiptera, Sternorrhyncha): a morphological and histological study.

Dorsal and ventral sides of the forewing of Aphis fabae (Scopoli 1763) (Sternorrhyncha, Hemiptera) were examined by scanning electron microscopy. Reinforcement elements on their surface as well as scale-like elements were described. Using histological methods, cross-sections of the material were done. They showed a two-layered membrane with a circular foramen inside. The course of veins and places of their bifurcation were followed. Common stem of radius (R), media (M), and cubitus anterior (CuA) veins were composed of separate tracheae, which ran separately at the beginning, then continued in a single tunnel, and finally disappeared. Nerves were not observed. Neither were tracheae visible on the further course of those veins. The presence of a devoid-of-trachea costal vein was confirmed. Under scanning electron microscope, convex reinforcements on dorsal side of the wing turned out to be more sclerotized parts of chitin, not giving a zigzag-like profile of the wing on sections. In this paper, we show for the first time a cross-section of a very delicate wing of an aphid representative.