Development of the ear, hearing capabilities and laterophysic connection in the spotfin butterflyfish (Chaetodon ocellatus)

The ontogeny of the ear, swim bladder and laterophysic connection was investigated in the spotfin butterflyfish, Chaetodon ocellatus in order to determine how the development of the laterophysic connection (a Chaetodon synapomorphy) is correlated with ontogenetic changes in the hearing capabilities in these abundant and ecologically important coral reef fishes. Histological and cleared and stained material revealed that the medial opening in the lateral line canal in the supracleithrum (which defines the laterophysic connection), an inflated physoclistous swim bladder, and the three otolithic organs are already present in the smallest individuals examined (7?C15?mm SL). The medial opening in the supracleithrum increases in size and the cylindrical swim bladder horns form after the loss of the head plates characteristic of the tholichthys stage, in individuals ??29?mm SL. The three sensory maculae of the ear increase in size, and the shape of the sacculus changes most dramatically with fish growth; hair cell density is highest in the utriculus. Physiological analysis of the reponse to sound pressure showed that larval and juvenile C. ocellatus had a hearing sensitivity peak at 100?C200?Hz, which was ~30?C40?dB more sensitive than that measured in larval coral reef fishes (e.g., damselfishes) that lack swim bladder horns. C. ocellatus did not show any ontogenetic changes in sensitivity to sound pressure, which may be explained by the fact that the growth of the swim bladder horns maintains the small distance between the swim bladder and ear that was established earlier during the larval stage. The timing of the development of the swim bladder horns suggests that if the laterophysic connection has a sensory acoustic function, its presence in individuals >29?mm SL suggests that its role is limited to post-settlement, reef-based behaviors.     

The laterophysic connection and swim bladder of butterflyfishes in the genus Chaetodon (Perciformes: Chaetodontidae)

The laterophysic connection (LC) is an association between bilaterally paired, anterior swim bladder extensions (horns) and medial openings in the supracleithral lateral line canals that diagnoses butterflyfishes in the genus Chaetodon. It has been hypothesized that the LC makes the lateral line system sensitive to sound pressure stimuli that are transmitted by the swim bladder horns and converted to fluid flow into the lateral line system via a laterophysic tympanum. The purpose of this study was to define variation in the morphology of the LC, swim bladder and swim bladder horns among 41 Chaetodon species from all 11 Chaetodon subgenera and a species from each of four non-Chaetodon genera using gross dissection, histological analysis as well as 2D or 3D CT (computed tomographic) imaging of live, anesthetized fishes. Our results demonstrate that the lateral line system appears rather unspecialized with well-ossified narrow canals in all species examined. Two LC types (direct and indirect), defined by whether or not the paired anterior swim bladder horns are in direct contact with a medial opening in the supracleithral lateral line canal, are found among species examined. Two variants on a direct LC and four variants of an indirect LC are defined by combinations of soft tissue anatomy (horn length [long/short] and width [wide/narrow], number of swim bladder chambers [one/two], and presence/absence of mucoid connective tissue in the medial opening in the supracleithrum). The combination of features defining each LC variant is predicted to have functional consequences for the bioacoustics of the system. These findings are consistent with the recent discovery that Chaetodon produce sounds during social interactions. The data presented here provide the comparative morphological context for the functional analysis of this novel swim bladder-lateral line connection.     

A proposed mechanism for the observed ontogenetic improvement in the hearing ability of hapuka (Polyprion oxygeneios)

Swim bladder extensions and hearing ability were examined in the temperate reef fish Polyprion oxygeneios (hapuka). Using the auditory evoked potential (AEP) technique, hearing thresholds were determined in four age-classes of hapuka, from larvae to juveniles. The youngest age-class had poor hearing abilities, with lowest thresholds of 132 dB re 1 μPa, and a narrow auditory bandwidth (100–800 Hz). Hearing ability improved significantly throughout the remainder of their first year, including decreases in thresholds of up to 27 dB, and an increase in auditory bandwidth (up to 1,000 Hz). Magnetic resonance imaging (MRI) was used to investigate structural mechanisms that may account for this ontogenetic improvement in hearing. These showed rostral extensions of the swim bladder developing early in the juvenile stage, and extending with increasing age closer to the otic capsule. It is suggested that this indirect connection between the swim bladder and the otic capsule could impart pressure sensitivity closer to the inner ear, accounting for the increase in sensitivity seen during development, although further investigation of older fish is required for conclusive evidence. The improvement in hearing ability in hapuka could be potentially related to a unique life history of extended pelagic durations up to 4 years.     

Swim bladder and posterior lateral line nerve of the nurseryfish, Kurtus gulliveri (Perciformes: Kurtidae)

The morphology of the swim bladder and inner ear of the nurseryfish, Kurtus gulliveri, appear adapted for enhanced pressure wave reception. The saccule is enlarged and surrounded by very thin bone and two large fontanelles that would present reduced resistance to pressure waves. The swim bladder is elaborate, with six dorsolaterally projecting pairs of lobes that are tightly encased in ribs and an additional caudally projecting pair of lobes encased in the first hemal spine. The ribs and musculature surrounding the swim bladder laterally are very thin, so that four or five "rib windows" are readily apparent on back-lit specimens. This swim bladder-rib configuration would also present reduced resistance to pressure waves to enhance function as a peripheral auditory structure. However, high-resolution X-ray computed tomography and dissection reveal no anterior projections of the swim bladder that could serve as a mechanical coupling to the inner ear. The posterior lateral line nerve is well developed and lies directly over the tips of the ribs encasing the swim bladder lobes. This nerve is not, however, associated with a lateral line canal and a lateral line canal is absent on most of the body. We hypothesize that the posterior lateral line nerve transmits mechanosensory information from the swim bladder.     

Relationship between Swim Bladder Morphology and Hearing Abilities-A Case Study on Asian and African Cichlids

Background

Several teleost species have evolved anterior extensions of the swim bladder which come close to or directly contact the inner ears. A few comparative studies have shown that these morphological specializations may enhance hearing abilities. This study investigates the diversity of swim bladder morphology in four Asian and African cichlid species and analyzes how this diversity affects their hearing sensitivity.

Methodology/Principal Findings

We studied swim bladder morphology by dissections and by making 3D reconstructions from high-resolution microCT scans. The auditory sensitivity was determined in terms of sound pressure levels (SPL) and particle acceleration levels (PAL) using the auditory evoked potential (AEP) recording technique. The swim bladders in Hemichromis guttatus and Steatocranus tinanti lacked anterior extensions and the swim bladder was considerably small in the latter species. In contrast, Paratilapia polleni and especially Etroplus maculatus possessed anterior extensions bringing the swim bladder close to the inner ears. All species were able to detect frequencies up to 3 kHz (SPL) except S. tinanti which only responded to frequencies up to 0.7 kHz. P. polleni and E. maculatus showed significantly higher auditory sensitivities at 0.5 and 1 kHz than the two species lacking anterior swim bladder extensions. The highest auditory sensitivities were found in E. maculatus, which possessed the most intimate swim bladder-inner ear relationship (maximum sensitivity 66 dB re 1 µPa at 0.5 kHz).

Conclusions

Our results indicate that anterior swim bladder extensions seem to improve mean absolute auditory sensitivities by 21–42 dB (SPLs) and 21–36 dB (PALs) between 0.5 and 1 kHz. Besides anterior extensions, the size of the swim bladder appears to be an important factor for extending the detectable frequency range (up to 3 kHz).     

Does the Hearing Sensitivity in Thorny Catfishes Depend on Swim Bladder Morphology?

Background

Thorny catfishes exhibit large variations in swim bladder morphology. These organs are of different sizes, forms and may have simple or branched diverticula. The swim bladder plays an important role in otophysans because it enhances their hearing sensitivity by transmitting sound pressure fluctuations via ossicles to the inner ear.

Methodology/Principal Findings

To investigate if a form-function relationship exists, the swim bladder morphology and hearing ability were analyzed in six species. The morphology was quantified by measuring the length, width and height and calculating a standardized swim bladder length (sSBL), which was then used to calculate the relative swim bladder length (rSBL). Hearing was measured using the auditory evoked potential (AEP) recording technique. Two species had simple apple-shaped and four species heart-shaped (cordiform) bladders. One of the latter species had short unbranched diverticula on the terminal margin, two had a secondary bladder and two had many long, branched diverticula. The rSBL differed significantly between most of the species. All species were able to detect frequencies between 70 Hz and 6 kHz, with lowest thresholds found between 0.5 and 1 kHz (60 dB re 1 µPa). Hearing curves were U-shaped except in Hemidoras morrisi in which it was ramp-like. Mean hearing thresholds of species possessing smaller rSBLs were slightly lower (maximum 8.5 dB) than those of species having larger rSBLs.

Conclusions/Significance

The current findings reveal a relationship between swim bladder form and its function among thorny catfishes. Relatively smaller swim bladders resulted in relatively better hearing. This is in contrast to a prior inter-familial study on catfishes in which species with large unpaired bladders possessed higher sensitivity at higher frequencies than species having tiny paired and encapsulated bladders.     

Intra‐ and Intersexual swim bladder dimorphisms in the plainfin midshipman fish (Porichthys notatus): Implications of swim bladder proximity to the inner ear for sound pressure detection

The plainfin midshipman fish, Porichthys notatus , is a nocturnal marine teleost that uses social acoustic signals for communication during the breeding season. Nesting type I males produce multiharmonic advertisement calls by contracting their swim bladder sonic muscles to attract females for courtship and spawning while subsequently attracting cuckholding type II males. Here, we report intra‐ and intersexual dimorphisms of the swim bladder in a vocal teleost fish and detail the swim bladder dimorphisms in the three sexual phenotypes (females, type I and II males) of plainfin midshipman fish. Micro‐computerized tomography revealed that females and type II males have prominent, horn‐like rostral swim bladder extensions that project toward the inner ear end organs (saccule, lagena, and utricle). The rostral swim bladder extensions were longer, and the distance between these swim bladder extensions and each inner‐ear end organ type was significantly shorter in both females and type II males compared to that in type I males. Our results revealed that the normalized swim bladder length of females and type II males was longer than that in type I males while there was no difference in normalized swim bladder width among the three sexual phenotypes. We predict that these intrasexual and intersexual differences in swim bladder morphology among midshipman sexual phenotypes will afford greater sound pressure sensitivity and higher frequency detection in females and type II males and facilitate the detection and localization of conspecifics in shallow water environments, like those in which midshipman breed and nest.     

3-D-orientation with the octavolateralis system.

Fish detect and localize a sound source with inner ear receptors and with the mechanosensory lateral line. The inner ear of fish is sensitive to the water displacements caused by sound waves through a direct, inertial response by hair cell epithelia of the ear. Hearing specialists, such as goldfish and herring, have accessory peripheral structures that provide additional sensitivity to the pressure component of a sound wave. While the inner ear of fish responds to the whole body motions caused by sound waves and--in case of hearing specialists--to sound pressure, the lateral line is only sensitive to water motions relative to the surface of the fish and to local pressure gradients. Using lateral line and/or acoustic input, some fish can determine the direction and the distance to a sound source. Most likely they do so by exploiting some of the mechanisms described in this paper. Piscivorous fish may use lateral line input to detect the wakes caused by swimming fish. Even in the absence of light catfish, for instance, can follow a 10 s old, three-dimensional wake left by a prey fish over distances up to 55 prey-body length.     

Hearing in the African lungfish (Protopterus annectens): pre-adaptation to pressure hearing in tetrapods?

Lungfishes are the closest living relatives of the tetrapods, and the ear of recent lungfishes resembles the tetrapod ear more than the ear of ray-finned fishes and is therefore of interest for understanding the evolution of hearing in the early tetrapods. The water-to-land transition resulted in major changes in the tetrapod ear associated with the detection of air-borne sound pressure, as evidenced by the late and independent origins of tympanic ears in all of the major tetrapod groups. To investigate lungfish pressure and vibration detection, we measured the sensitivity and frequency responses of five West African lungfish (Protopterus annectens) using brainstem potentials evoked by calibrated sound and vibration stimuli in air and water. We find that the lungfish ear has good low-frequency vibration sensitivity, like recent amphibians, but poor sensitivity to air-borne sound. The skull shows measurable vibrations above 100 Hz when stimulated by air-borne sound, but the ear is apparently insensitive at these frequencies, suggesting that the lungfish ear is neither adapted nor pre-adapted for aerial hearing. Thus, if the lungfish ear is a model of the ear of early tetrapods, their auditory sensitivity was limited to very low frequencies on land, mostly mediated by substrate-borne vibrations.     

Effects of exposure to pile-driving sounds on the lake sturgeon,Nile tilapia and hogchoker

Pile-driving and other impulsive sound sources have the potential to injure or kill fishes. One mechanism that produces injuries is the rapid motion of the walls of the swim bladder as it repeatedly contacts nearby tissues. To further understand the involvement of the swim bladder in tissue damage, a specially designed wave tube was used to expose three species to pile-driving sounds. Species included lake sturgeon (Acipenser fulvescens)—with an open (physostomous) swim bladder, Nile tilapia (Oreochromis niloticus)—with a closed (physoclistous) swim bladder and the hogchoker (Trinectes maculatus)—a flatfish without a swim bladder. There were no visible injuries in any of the exposed hogchokers, whereas a variety of injuries were observed in the lake sturgeon and Nile tilapia. At the loudest cumulative and single-strike sound exposure levels (SEL_cum and SEL_ss respectively), the Nile tilapia had the highest total injuries and the most severe injuries per fish. As exposure levels decreased, the number and severity of injuries were more similar between the two species. These results suggest that the presence and type of swim bladder correlated with injury at higher sound levels, while the extent of injury at lower sound levels was similar for both kinds of swim bladders.     

Better than fish on land? Hearing across metamorphosis in salamanders

Early tetrapods faced an auditory challenge from the impedance mismatch between air and tissue in the transition from aquatic to terrestrial lifestyles during the Early Carboniferous (350 Ma). Consequently, tetrapods may have been deaf to airborne sounds for up to 100 Myr until tympanic middle ears evolved during the Triassic. The middle ear morphology of recent urodeles is similar to that of early ‘lepospondyl’ microsaur tetrapods, and experimental studies on their hearing capabilities are therefore useful to understand the evolutionary and functional drivers behind the shift from aquatic to aerial hearing in early tetrapods. Here, we combine imaging techniques with neurophysiological measurements to resolve how the change from aquatic larvae to terrestrial adult affects the ear morphology and sensory capabilities of salamanders. We show that air-induced pressure detection enhances underwater hearing sensitivity of salamanders at frequencies above 120 Hz, and that both terrestrial adults and fully aquatic juvenile salamanders can detect airborne sound. Collectively, these findings suggest that early atympanic tetrapods may have been pre-equipped to aerial hearing and are able to hear airborne sound better than fish on land. When selected for, this rudimentary hearing could have led to the evolution of tympanic middle ears.     

Hearing in Cichlid Fishes under Noise Conditions

Background

Hearing thresholds of fishes are typically acquired under laboratory conditions. This does not reflect the situation in natural habitats, where ambient noise may mask their hearing sensitivities. In the current study we investigate hearing in terms of sound pressure (SPL) and particle acceleration levels (PAL) of two cichlid species within the naturally occurring range of noise levels. This enabled us to determine whether species with and without hearing specializations are differently affected by noise.

Methodology/Principal Findings

We investigated auditory sensitivities in the orange chromide Etroplus maculatus, which possesses anterior swim bladder extensions, and the slender lionhead cichlid Steatocranus tinanti, in which the swim bladder is much smaller and lacks extensions. E. maculatus was tested between 0.2 and 3kHz and S. tinanti between 0.1 and 0.5 kHz using the auditory evoked potential (AEP) recording technique. In both species, SPL and PAL audiograms were determined in the presence of quiet laboratory conditions (baseline) and continuous white noise of 110 and 130 dB RMS. Baseline thresholds showed greatest hearing sensitivity around 0.5 kHz (SPL) and 0.2 kHz (PAL) in E. maculatus and 0.2 kHz in S. tinanti. White noise of 110 dB elevated the thresholds by 0–11 dB (SPL) and 7–11 dB (PAL) in E. maculatus and by 1–2 dB (SPL) and by 1–4 dB (PAL) in S. tinanti. White noise of 130 dB elevated hearing thresholds by 13–29 dB (SPL) and 26–32 dB (PAL) in E. maculatus and 6–16 dB (SPL) and 6–19 dB (PAL) in S. tinanti.

Conclusions

Our data showed for the first time for SPL and PAL thresholds that the specialized species was masked by different noise regimes at almost all frequencies, whereas the non-specialized species was much less affected. This indicates that noise can limit sound detection and acoustic orientation differently within a single fish family.     

Acoustic overstimulation activates 5'-AMP-activated protein kinase through a temporary decrease in ATP level in the cochlear spiral ligament prior to permanent hearing loss in mice

Inner ear disorders are known to be elicited by mitochondrial dysfunction, which decreases the ATP level in the inner ear. 5′-AMP-activated protein kinase (AMPK) is a serine/threonine kinase activated by metabolic stress and by an increase in the AMP/ATP ratio. To elucidate the involvement of AMPK-derived signals in noise-induced hearing loss, we investigated whether in vivo acoustic overstimulation would activate AMPK in the cochlea of mice. Std-ddY mice were exposed to 8 kHz octave band noise at a 90-, 110- or 120-dB sound pressure level (SPL) for 2 h. Exposure to the noise at 110 or 120 dB SPL produced outer hair cell death in the organ of Corti and permanent hearing loss. Exposure to the noise at 120-dB SPL elevated the level of the phospho-AMPK α-subunit (p-AMPKα), without affecting the protein level of this subunit, immediately and at 12-h post-exposure in the lateral wall structures including the spiral ligament and stria vascularis. In the hair cells and spiral ganglion cells, no marked change in the level of p-AMPKα was observed at any time post-exposure. The level of phospho-c-Jun N-terminal kinase (p-JNK) was increased in the lateral wall structures at 2- to 4-h post-exposure at 120 dB SPL. Noise exposure significantly, but temporarily, decreased the ATP level in the spiral ligament, in an SPL-dependent manner at 110 dB and above. Likewise, elevation of p-AMPKα and p-JNK levels was also observed in the lateral wall structures post-exposure to noise at an SPL of 110 dB and above. Taken together, our data suggest that AMPK and JNK were activated by ATP depletion in the cochlear spiral ligament prior to permanent hearing loss induced by in vivo acoustic overstimulation.     

Morphological development of the swim bladder in hatchery-reared striped trumpeter Latris lineata

This study examined swim bladder morphogenesis in three cohorts of striped trumpeter (Latris lineata), a euphysoclist species with physostomous larvae. The swim bladder was first discernible 1–2 days after hatching as an evagination on the dorsal surface of the incipient digestive tract. It comprised a cluster of mesenchymal cells surrounding an inner primordium of epithelial cells. At mouth opening in larvae of 5.3 mm standard length (SL), the swim bladder was noticeably enlarged. Histologically, the swim bladder lumen was dilated and liquid filled. The pneumatic duct was first seen during the dilation stage and the rete mirabile began forming among the connective tissue surrounding the swim bladder. Initial swim bladder inflation occurred on day 11 post‐hatching in Cohort 1, at 14°C, and day 9 post‐hatching, in Cohorts 2 and 3, at 16°C. Histologically, the lumens of inflated swim bladders were ellipsoid and the epithelium was squamous, except for cuboidal gas gland cells at the anterio‐ventral and anterio‐lateral regions of the swim bladder. During the initial inflation interval the pneumatic duct was dilated in larvae both with and without swim bladder inflation. The pneumatic duct began to regress in some larvae over 7.5 mm SL. The swim bladder of striped trumpeter was similar to larvae of other altricial perciform marine fish in respect to organ derivation, tissue differentiation, luminal dilation and initial gaseous inflation. However, variations, particularly the delay in initial swim bladder inflation until after the start of feeding, were observed that could be fundamental to problems encountered during larval rearing.     

Hyperacute directional hearing and phonotactic steering in the cricket (Gryllus bimaculatus deGeer)

Background

Auditory mate or prey localisation is central to the lifestyle of many animals and requires precise directional hearing. However, when the incident angle of sound approaches 0° azimuth, interaural time and intensity differences gradually vanish. This poses a demanding challenge to animals especially when interaural distances are small. To cope with these limitations imposed by the laws of acoustics, crickets employ a frequency tuned peripheral hearing system. Although this enhances auditory directionality the actual precision of directional hearing and phonotactic steering has never been studied in the behaviourally important frontal range.

Principal Findings

Here we analysed the directionality of phonotaxis in female crickets (Gryllus bimaculatus) walking on an open-loop trackball system by measuring their steering accuracy towards male calling song presented at frontal angles of incidence. Within the range of ±30°, females reliably discriminated the side of acoustic stimulation, even when the sound source deviated by only 1° from the animal''s length axis. Moreover, for angles of sound incidence between 1° and 6° the females precisely walked towards the sound source. Measuring the tympanic membrane oscillations of the front leg ears with a laser vibrometer revealed between 0° and 30° a linear increasing function of interaural amplitude differences with a slope of 0.4 dB/°. Auditory nerve recordings closely reflected these bilateral differences in afferent response latency and intensity that provide the physiological basis for precise auditory steering.

Conclusions

Our experiments demonstrate that an insect hearing system based on a frequency-tuned pressure difference receiver achieves directional hyperacuity which easily rivals best directional hearing in mammals and birds. Moreover, this directional accuracy of the cricket''s hearing system is reflected in the animal''s phonotactic motor response.     

Listening to the environment: hearing differences from an epigenetic effect in solitarious and gregarious locusts

Locusts display a striking form of phenotypic plasticity, developing into either a lone-living solitarious phase or a swarming gregarious phase depending on population density. The two phases differ extensively in appearance, behaviour and physiology. We found that solitarious and gregarious locusts have clear differences in their hearing, both in their tympanal and neuronal responses. We identified significant differences in the shape of the tympana that may be responsible for the variations in hearing between locust phases. We measured the nanometre mechanical responses of the ear''s tympanal membrane to sound, finding that solitarious animals exhibit greater displacement. Finally, neural experiments signified that solitarious locusts have a relatively stronger response to high frequencies. The enhanced response to high-frequency sounds in the nocturnally flying solitarious locusts suggests greater investment in detecting the ultrasonic echolocation calls of bats, to which they are more vulnerable than diurnally active gregarious locusts. This study highlights the importance of epigenetic effects set forth during development and begins to identify how animals are equipped to match their immediate environmental needs.     

Dynamics Modeling and Simulation of a Bionic Swim Bladder System in Underwater Robotics

While considering that fish could suspend themselves under water and could enhance their mobility by adjusting its swim bladder, we have carried out research on a bionic swim bladder system in underwater robotics, which could amend the underwater robotics' static balance and controllability conditions even if the depth of water changes. First, this paper introduces the bio-swim bladder's structure and function. Second, it works out the dynamic model of the bionic swim bladder, and then it analyses the dynamic characteristic and effect of the bionic swim bladder system with the software Matlab/simulink. Finally, considering about the nonlinear relationship of the parameters in the model, this paper brings forward a dual-speed control method, which could make the effect of the bionic swim bladder non-coupling. The result of the simulation reveals that the bionic swim bladder could change the buoyancy and centroid distribution of the underwater robotics effectively and independently, bringing it into a balance state, under which the control and maneuverability could be enhanced.     

Bone‐conduction hearing and seismic sensitivity of the late permian anomodont Kawingasaurus fossilis

An investigation of the internal cranial anatomy of the anomodont Kawingasaurus from the Upper Permian Usili Formation in Tanzania by means of neutron tomography revealed an unusual inner and middle ear anatomy such as extraordinarily inflated vestibules, lateroventrally orientated stapes with large footplates, and a small angle between the planes of the anterior and lateral semicircular canals. The vestibule has a volume, which is about 25 times larger than the human vestibule, although Kawingasaurus has only a skull length of approximately 40 mm. Vestibule inflation and enlarged stapes footplates are thought to be functionally correlated with bone‐conduction hearing; both morphologies have been observed in fossorial vertebrates using seismic signals for communication. The firmly fused triangular head with spatulate snout was probably used for digging and preadapted to seismic signal detection. The quadrate‐quadratojugal complex was able to transmit sound from the articular to the stapes by small vibrations of the quadrate process, which formed a ball and socket joint with the squamosal. Mechanical considerations suggest that the ventrolaterally orientated stapes of Kawingasaurus was mechanically better suited to transmit seismic sound from the ground to the fenestra vestibuli than a horizontal orientated stapes. The low sound pressure level transformer ratio of 2–3 in Kawingasaurus points to a seismic sensitivity of the middle ear and a vestigial or reduced sensitivity to airborne sound. Three hypothetical pathways of bone conduction in Kawingasaurus are discussed: 1) sound transmission via the spatulate snout and skull roof to the otic capsules, 2) relative movements resulting from the inertia of the mandible if sound is percepted with the skull, and 3) bone conduction from the substrate via mandible, jaw articulation, and stapes to the inner ear. J. Morphol. 276:121–143, 2015. © 2014 Wiley Periodicals, Inc.     

昆虫弦音器及其声音感受分子机制

弦音器是昆虫类特有的一种机械感受器,亦称弦音感受器或剑梢感受器。它主要具有感知外界声压和体内肌肉运动的听觉功能,研究弦音器的机能结构对揭秘昆虫听觉的神经机制有重要的科学意义。本文从弦音器多样性和进化入手,重点综述了弦音器的微细结构、基因功能定位、声音感受分子机制及其声压增幅分子生物物理学原理,为昆虫听觉仿生学的研究提供了理论依据。     

Characterization of the primary sonic muscles in Carapus acus (Carapidae): a multidisciplinary approach

Sound production in carapid fishes results from the action of extrinsic muscles that insert into the swim bladder. Biochemical, histochemical and morphological techniques were used to examine the sonic muscles and compare them with epaxial muscles in Carapus acus. Sonic fibres are thicker than red and thinner than white epaxial fibres, and sonic fibres and myofibrils exhibit an unusual helicoidal organization: the myofibrils of the centre are in a straight line whereas they are more and more twisted towards the periphery. Sonic muscles have both features of red (numerous mitochondria, high glycogen content) and white (alkali-stable ATPase) fibres. They differ also in the isoforms of the light chain (LC3) and heavy chain (HC), in having T tubules at both the Z-line and the A-I junction and in a unique parvalbumin isoform (PAI) that may aid relaxation. All these features lead to the expression of two assumptions about sound generation: the sonic muscle should be able to perform fast and powerful contractions that provoke the forward movement of the forepart of the swim bladder and the stretching and "flapping" of the swim bladder fenestra; the helicoidal organization allows progressive drawing of the swim bladder fenestra which emits a sound when rapidly released in a spring-like manner.