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Male crickets attract females by producing calls with their forewings. Louder calls travel further and are more effective at attracting mates. However, crickets are small, and therefore inefficient, dipole sound-sources. Only a small group called tree crickets make acoustic tools called baffles which reduce acoustic short-circuiting, a source of dipole inefficiency. Here, we ask why baffling is uncommon among crickets. We hypothesize that baffling may be rare, because like other tools they offer insufficient advantage for most species. To test this, we modelled the calling efficiencies of crickets within the full space of possible natural wing sizes and call frequencies, in multiple acoustic environments. We then generated efficiency landscapes, within which we plotted the positions of 111 cricket species across 7 phylogenetic clades. We found that all sampled crickets, in all conditions, could gain efficiency from tool use. Surprisingly, however, we also found that calling from the ground significantly increased efficiency, with or without a baffle, by as much as an order of magnitude. We found that the ground provides some reduction of acoustic short-circuiting but also halves the air volume within which sound is radiated. It simultaneously reflects sound upwards, allowing recapture of a significant amount of acoustic energy through constructive interference. Thus, using the ground as a reflective baffle is a very effective strategy for increasing calling efficiency. Indeed, theory suggests that this increase in efficiency is accessible not just to crickets, but to all acoustically communicating animals whether they are dipole or monopole sound sources.Significance StatementLoudness is a crucial functional feature of calls in acoustically communicating animals. Animals attempting to reach prospective mates or ward off predators are expected to make themselves as loud as possible. Given the importance of loudness, there are two long-standing, seemingly unrelated paradoxes in acoustic communication. The first is the rarity of acoustic tool use. The second is the high number of animals that call from reflective surfaces, like the ground, known be an impediment to sound propagation. We resolve both paradoxes and show them to be related. By refocusing analysis from sound propagation to sound radiation, we show that the ground is the opposite of an impediment and can, in fact, boost sound loudness more than tool use. Thus, we show that calling from a reflective surface is an alternative strategy for maximizing call loudness, and one that is available to all animals.

Bush-crickets (or katydids) have sophisticated and ultrasonic ears located in the tibia of their forelegs, with a working mechanism analogous to the mammalian auditory system. Their inner-ears are endowed with an easily accessible hearing organ, thecrista acustica(CA), possessing a spatial organisation that allows for different frequencies to be processed at specific graded locations within the structure. Similar to the basilar membrane in the mammalian ear, the CA contains mechanosensory receptors which are activated through the frequency dependent displacement of the CA. While this tonotopical arrangement is generally attributed to the gradual stiffness and mass changes along the hearing organ, the mechanisms behind it have not been analysed in detail. In this study, we take a numerical approach to investigate this mechanism in theCopiphora gorgonensisear. In addition, we propose and test the effect of the different vibration transmission mechanisms on the displacement of the CA. The investigation was carried out by conducting finite-element analysis on a three-dimensional, idealised geometry of theC. gorgonensisinner-ear, which was based on precise measurements. The numerical results suggested that(i)even the mildest assumptions about stiffness and mass gradients allow for tonotopy to emerge, and(ii)the loading area and location for the transmission of the acoustic vibrations play a major role in the formation of tonotopy.

Our ability to sense and move our bodies relies on proprioceptors, sensory neurons that detect mechanical forces within the body. Proprioceptors are diverse: different subtypes detect different features of joint kinematics, such as position, directional movement, and vibration. However, because they are located within complex and dynamic peripheral tissues, the underlying mechanisms of proprioceptor feature selectivity remain poorly understood. Here, we investigate molecular and biomechanical contributions to proprioceptor diversity in theDrosophilaleg. Using single-nucleus RNA sequencing, we found that different proprioceptor subtypes express similar complements of mechanosensory and other ion channels. However, anatomical reconstruction of the proprioceptive organ and connected tendons revealed major biomechanical differences between proprioceptor subtypes. We constructed a computational model of the proprioceptors and tendons, which identified a putative biomechanical mechanism for joint angle selectivity. The model also predicted the existence of a goniotopic map of joint angle among position-tuned proprioceptors, which we confirmed using calcium imaging. Our findings suggest that biomechanical specialization is a key determinant of proprioceptor feature selectivity inDrosophila. More broadly, our discovery of proprioceptive maps in the fly leg reveals common organizational principles between proprioception and other topographically organized sensory systems.

Jumping in animals presents an interesting locomotory strategy as it requires the generation of large forces and accurate timing. Jumping in arachnids is further complicated by their semi-hydraulic locomotion system. Among arachnids, jumping spiders (Family Salticidae) are agile and dexterous jumpers. However, less is known about jumping in small salticid species. Here we used Habronattus conjunctus, a small jumping spider (body length ~ 4.5 mm) to examine its jumping performance and compare it to that of other jumping spiders and insects. We also explored how legs are used during the takeoff phase of jumps. Jumps were staged between two raised platforms. We analyzed jumping videos with DeepLabCut to track 21 points on the cephalothorax, abdomen, and legs. By analyzing leg liftoff and extension patterns, we found evidence that H. conjunctus primarily uses the third legs to power jumps. We also found that H. conjunctus jumps achieve lower takeoff speeds and accelerations than most other jumping arthropods, including other jumping spiders. Habronattus conjunctus takeoff time was similar to other jumping arthropods of the same body mass. We discuss the mechanical benefits and drawbacks of a semi-hydraulic system of locomotion and consider how small spiders may extract dexterous jumps from this locomotor system.

Motivated by recent developments suggesting that interaural coupling in non-mammals allows for the two active ears to effectively synchronize, this report describes otoacoustic measurements made in the oral cavity of lizards. As expected from that model, spontaneous otoacoustic emissions (SOAEs) were readily measurable in the mouth, which is contiguous with the interaural airspace. Additionally, finite element model calculations were made to simulate the interaural acoustics based upon SOAE-related tympanic membrane vibrational data. Taken together, these data support the notion of two active ears synchronizing by virtue of acoustic coupling and have potential implications for sound localization at low-levels.

Jumping in animals presents an interesting locomotory strategy as it requires the generation of large forces and accurate timing. Jumping in arachnids is further complicated by their semi-hydraulic locomotion system. Among arachnids, jumping spiders (Family Salticidae) are agile and dexterous jumpers. However, less is known about jumping in small salticid species. Here we used Habronattus conjunctus, a small jumping spider (body length ~ 4.5 mm) to examine its jumping performance and compare it to that of other jumping spiders and insects. We also explored how legs are used during the takeoff phase of jumps. Jumps were staged between two raised platforms. We analyzed jumping videos with DeepLabCut to track 21 points on the cephalothorax, abdomen, and legs. By analyzing leg liftoff and extension patterns, we found evidence that H. conjunctus primarily uses the third legs to power jumps. We also found that H. conjunctus jumps achieve lower takeoff speeds and accelerations than most other jumping arthropods, including other jumping spiders. Habronattus conjunctus takeoff time was similar to other jumping arthropods of the same body mass. We discuss the mechanical benefits and drawbacks of a semi-hydraulic system of locomotion and consider how small spiders may extract dexterous jumps from this locomotor system.

Khoomei is a unique singing style originating from the republic of Tuva in central Asia. Singers produce two pitches simultaneously: a booming low-frequency rumble alongside a hovering high-pitched whistle-like tone. The biomechanics of this biphonation are not well-understood. Here, we use sound analysis, dynamic magnetic resonance imaging, and vocal tract modeling to demonstrate how biphonation is achieved by modulating vocal tract morphology. Tuvan singers show remarkable control in shaping their vocal tract to narrowly focus the harmonics (or overtones) emanating from their vocal cords. The biphonic sound is a combination of the fundamental pitch and a focused filter state, which is at the higher pitch (1–2 kHz) and formed by merging two formants, thereby greatly enhancing sound-production in a very narrow frequency range. Most importantly, we demonstrate that this biphonation is a phenomenon arising from linear filtering rather than from a nonlinear source., eLife digest The republic of Tuva, a remote territory in southern Russia located on the border with Mongolia, is perhaps best known for its vast mountainous geography and the unique cultural practice of “throat singing”. These singers simultaneously create two different pitches: a low-pitched drone, along with a hovering whistle above it. This practice has deep cultural roots and has now been shared more broadly via world music performances and the 1999 documentary Genghis Blues. Despite many scientists being fascinated by throat singing, it was unclear precisely how throat singers could create two unique pitches. Singing and speaking in general involves making sounds by vibrating the vocal cords found deep in the throat, and then shaping those sounds with the tongue, teeth and lips as they move up the vocal tract and out of the body. Previous studies using static images taken with magnetic resonance imaging (MRI) suggested how Tuvan singers might produce the two pitches, but a mechanistic understanding of throat singing was far from complete. Now, Bergevin et al. have better pinpointed how throat singers can produce their unique sound. The analysis involved high quality audio recordings of three Tuvan singers and dynamic MRI recordings of the movements of one of those singers. The images showed changes in the singer’s vocal tract as they sang inside an MRI scanner, providing key information needed to create a computer model of the process. This approach revealed that Tuvan singers can create two pitches simultaneously by forming precise constrictions in their vocal tract. One key constriction occurs when tip of the tongue nearly touches a ridge on the roof of the mouth, and a second constriction is formed by the base of the tongue. The computer model helped explain that these two constrictions produce the distinctive sounds of throat singing by selectively amplifying a narrow set of high frequency notes that are made by the vocal cords. Together these discoveries show how very small, targeted movements of the tongue can produce distinctive sounds.

The Drosophila auditory system consists of four large basal segments: the arista, the funiculus, the pedicel, and the scape. When an acoustic stimulus is applied to the arista and the funiculus their mechanical vibrations are transmitted to chordotonal neurons in Johnston’s organ where mechanoelectric transduction arises. We study the mechanotransduction mechanism in the Drosophila auditory system by using a laser Doppler vibrometer (LDV) and extracellular electrophysiology. We find that large and small peaks appear alternatively and that the antenna vibration is asymmetric depending on whether the pedicel and the scape are fixed. Interestingly, we find that this asymmetric vibration accompanies the alternating neural peak structure. Here, we propose a mathematical model to explain the alternating peak structure by using a model consisting of two opposing neurons that are modeled as strings. Generally, strings have tension only when they are elongated. This property allows the alternating neural peaks for asymmetric antenna motion.

Tree crickets use sound to attract mates and make acoustic baffles to increase their sound production efficiency. It has been recently discovered that these insects use a flexible yet inherited behavioural programme to make acoustically optimal baffles. Whether these baffles qualify as tools, however, remains controversial. Here, baffle‐using and baffle‐making behaviours are analysed using the most current and authoritative definition of tool use. The current definition of tool use does not require the tool to be detached from the substrate and includes attached but manipulable external objects. Given this schema, tree cricket baffles, which are attached but manipulated prior to use, must be considered tools. The mode of manufacture for a baffle is “Subtract,” and the mode of use is “Drape”.

Khoomei is a unique singing style originating from the Central Asian republic of Tuva. Singers produce two pitches simultaneously: a booming low-frequency rumble alongside a hovering high-pitched whistle-like tone. The biomechanics of this biphonation are not well-understood. Here, we use sound analysis, dynamic magnetic resonance imaging, and vocal tract modeling to demonstrate how biphonation is achieved by modulating vocal tract morphology. Tuvan singers show remarkable control in shaping their vocal tract to narrowly focus the harmonics (or overtones) emanating from their vocal cords. The biphonic sound is a combination of the fundamental pitch and a focused filter state, which is at the higher pitch (1-2 kHz) and formed by merging two formants, thereby greatly enhancing sound-production in a very narrow frequency range. Most importantly, we demonstrate that this biphonation is a phenomenon arising from linear filtering rather than a nonlinear source.

Spiders rely on mechanical vibration sensing for sexual signalling, prey capture and predator evasion. The sensory organs underlying vibration detection, called slit sensilla, resemble cracks in the spider's exoskeleton, and are distributed all over the spider body. Those crucial to sensing web- and other substrate-borne vibrations are called lyriform organs and are densely distributed around leg joints. It has been shown that forces that cause bending at leg joints also activate these lyriform organs. Little is known of how the biomechanics of the body of a freely-suspended spider in its natural posture interact with vibrations introduced into the body and how this affects vibration perception. Female black widow spiders, in particular, have a striking body-form; their long thin legs support a large pendulous abdomen. Here, we show that in their natural posture, the large abdominal mass of black widow females, interacts with the spring-like behaviour of their leg joints and determines the mechanical behaviour of different leg joints. Furthermore, we find that adopting different body postures enables females to alter both the level and tuning of the mechanical input to lyriform organs. Therefore, we suggest that posture may be used to flexibly and reversibly focus attention to different classes or components of web vibration. Postural effects thus emphasize the dynamic loop of interactions between behaviour and perception, i.e. between 'brain' and body.

Insects have evolved a diversity of hearing organs specialized to detect sounds critical for survival. We report on a unique structure on butterfly wings that enhances hearing. The Satyrini are a diverse group of butterflies occurring throughout the world. One of their distinguishing features is a conspicuous swelling of their forewing vein, but the functional significance of this structure is unknown. Here, we show that wing vein inflations function in hearing. Using the common wood nymph, Cercyonis pegala , as a model, we show that (i) these butterflies have ears on their forewings that are most sensitive to low frequency sounds (less than 5 kHz); (ii) inflated wing veins are directly connected to the ears; and (iii) when vein inflations are ablated, sensitivity to low frequency sounds is impaired. We propose that inflated veins contribute to low frequency hearing by impedance matching.

Generated inside the inner ear, spontaneous otoacoustic emissions (SOAEs) are the most salient evidence for an “active” ear. These emissions propagate back through the middle and external ears, setting the tympanic membrane in motion such that it effectively acts as a “speaker”. Better characterization of this motion would be useful for the quantification of power produced by the associated active processes. Towards this end, we took a comparative approach, examining spontaneous activity in both a lizard (green anole, Anolis carolinensis) and an invertebrate (tree cricket, Oecanthus nigricornis). Using a laser Doppler vibrometer and sensitive OAE probe, we measured both tympanum motion and the acoustic output. For lizards, using a single point measurement system (whose focal point was not well localized), roughly similar behavior was observed in both the mechanical tympanum motion and the acoustic SOAE. For a relatively large spontaneous vibration peak (at 3.9 kHz) on the lizard tympanum, the amplitude of displacement was approximately 60 pm. Differences were however noted between SOAE and vibration, and several possible explanations for such are discussed. For the cricket, only “open” coupling of the microphone could be implemented. It was observed that while spontaneous vibration on the tympanum could be detected above the noise floor (single peak about 3-4 kHz, with displacements of 10-35 pm), SOAEs could not. When a stimulus was presented to a elicit a stimulus frequency emission (SFOAE) using a suppression paradigm, no residual could be detected either in the vibration or acoustic response. Lastly, when two tones were presented, distortions were readily observable in the tympanum motion but no acoustic DPOAEs were detected. It is suggested that the relatively small size of the cricket tympanum and open coupling employed could explain these inconsistencies (e.g., it is too small to effectively radiate acoustic power above the noise floor).Generated inside the inner ear, spontaneous otoacoustic emissions (SOAEs) are the most salient evidence for an “active” ear. These emissions propagate back through the middle and external ears, setting the tympanic membrane in motion such that it effectively acts as a “speaker”. Better characterization of this motion would be useful for the quantification of power produced by the associated active processes. Towards this end, we took a comparative approach, examining spontaneous activity in both a lizard (green anole, Anolis carolinensis) and an invertebrate (tree cricket, Oecanthus nigricornis). Using a laser Doppler vibrometer and sensitive OAE probe, we measured both tympanum motion and the acoustic output. For lizards, using a single point measurement system (whose focal point was not well localized), roughly similar behavior was observed in both the mechanical tympanum motion and the acoustic SOAE. For a relatively large spontaneous vibration peak (at 3.9 kHz) on the lizard tympanum, the amplitude of...

Tree cricket hearing shows all the features of an actively amplified auditory system, particularly spontaneous oscillations (SOs) of the tympanal membrane. As expected from an actively amplified auditory system, SO frequency and the peak frequency in evoked responses as observed in sensitivity spectra are correlated. Sensitivity spectra also show compressive non-linearity at this frequency, i.e. a reduction in peak height and sharpness with increasing stimulus amplitude. Both SO and amplified frequency also change with ambient temperature, allowing the auditory system to maintain a filter that is matched to song frequency. In tree crickets, remarkably, song frequency varies with ambient temperature. Interestingly, active amplification has been reported to be switched ON and OFF. The mechanism of this switch is as yet unknown. In order to gain insights into this switch, we recorded and analysed SOs as the auditory system transitioned from the passive (OFF) state to the active (ON) state. We found that while SO amplitude did not follow a fixed pattern, SO frequency changed during the ON-OFF transition. SOs were first detected above noise levels at low frequencies, sometimes well below the known song frequency range (0.5-1 kHz lower). SO frequency was observed to increase over the next ∼30 minutes, in the absence of any ambient temperature change, before settling at a frequency within the range of conspecific song. We examine the frequency shift in SO spectra with temperature and during the ON/OFF transition and discuss the mechanistic implications. To our knowledge, such modulation of active auditory amplification, and its dynamics are unique amongst auditory animals.Tree cricket hearing shows all the features of an actively amplified auditory system, particularly spontaneous oscillations (SOs) of the tympanal membrane. As expected from an actively amplified auditory system, SO frequency and the peak frequency in evoked responses as observed in sensitivity spectra are correlated. Sensitivity spectra also show compressive non-linearity at this frequency, i.e. a reduction in peak height and sharpness with increasing stimulus amplitude. Both SO and amplified frequency also change with ambient temperature, allowing the auditory system to maintain a filter that is matched to song frequency. In tree crickets, remarkably, song frequency varies with ambient temperature. Interestingly, active amplification has been reported to be switched ON and OFF. The mechanism of this switch is as yet unknown. In order to gain insights into this switch, we recorded and analysed SOs as the auditory system transitioned from the passive (OFF) state to the active (ON) state. We found that whil...

1. The subtitle Laser Doppler vibrometer measurement in “EXPERIMENTS AND DISCUSSION” should be replaced by PHYSICAL MODEL FOR DROSOPHILA ANTENNA” 2. The first paragraph of “EXPERIMENTS AND DISCUSSION” deals with “Laser Doppler vibrometer measurement”.

Tree cricket males produce tonal songs, used for mate-attraction and male-male interactions. Active mechanics tunes hearing to conspecific song frequency. However, tree cricket song frequency increases with temperature, presenting a problem for tuned listeners. We show that the actively amplified frequency increases with temperature, thus shifting mechanical and neuronal auditory tuning to maintain a match with conspecific song frequency. Active auditory processes are known from several taxa, but their adaptive function has rarely been demonstrated. We show that tree crickets harness active processes to ensure that auditory tuning remains matched to conspecific song frequency, despite changing environmental conditions and signal characteristics. Adaptive tuning allows tree crickets to selectively detect potential mates or rivals over large distances and is likely to bestow a strong selective advantage by reducing mate-finding effort and facilitating intermale interactions.

Tree cricket males produce tonal songs, which are used to attract mates and to interact with other males. In other crickets, the mechanical properties of their peripheral auditory system have evolved to have resonant modes at the communication frequency. The tree cricket auditory system, however, is not passively tuned to song frequency. Tree crickets exploit an active amplification process to tune hearing to song frequency. However, their song frequency increases with temperature, presenting a problem for tuned listeners. We show here that the actively amplified frequency increases with temperature, and shifts mechanical and neuronal auditory tuning to maintain a match with changing song frequency. We also find that in tree crickets active amplification does not provide greater sensitivity or dynamic range than that observed in other crickets that lack active amplification. Thus, the primary adaptive function of active amplification is to ensure that auditory tuning remains matched to conspecific song frequency, despite changing environmental conditions and signal characteristics.Tree cricket males produce tonal songs, which are used to attract mates and to interact with other males. In other crickets, the mechanical properties of their peripheral auditory system have evolved to have resonant modes at the communication frequency. The tree cricket auditory system, however, is not passively tuned to song frequency. Tree crickets exploit an active amplification process to tune hearing to song frequency. However, their song frequency increases with temperature, presenting a problem for tuned listeners. We show here that the actively amplified frequency increases with temperature, and shifts mechanical and neuronal auditory tuning to maintain a match with changing song frequency. We also find that in tree crickets active amplification does not provide greater sensitivity or dynamic range than that observed in other crickets that lack active amplification. Thus, the primary adaptive function of active amplification is to ensure that auditory tuning remains matched to conspecific song fr...

Crickets have two tympanal membranes on the tibiae of each foreleg. Among several field cricket species of the genus Gryllus (Gryllinae), the posterior tympanal membrane (PTM) is significantly larger than the anterior membrane (ATM). Laser Doppler vibrometric measurements have shown that the smaller ATM does not respond as much as the PTM to sound. Hence the PTM has been suggested to be the principal tympanal acoustic input to the auditory organ. In tree crickets (Oecanthinae), the ATM is slightly larger than the PTM. Both membranes are structurally complex, presenting a series of transverse folds on their surface, which are more pronounced on the ATM than on the PTM. The mechanical response of both membranes to acoustic stimulation was investigated using microscanning laser Doppler vibrometry. Only a small portion of the membrane surface deflects in response to sound. Both membranes exhibit similar frequency responses, and move out of phase with each other, producing compressions and rarefactions of the tracheal volume backing the tympanum. Therefore, unlike field crickets, tree crickets may have four instead of two functional tympanal membranes. This is interesting in the context of the outstanding question of the role of spiracular inputs in the auditory system of tree crickets.

SUMMARY Animals have to accomplish several tasks in their lifetime, such as finding food and mates and avoiding predators. Animals that locate these using sound need to detect, recognize and localize appropriate acoustic objects in their environment, typically in noisy, non-ideal conditions. Quantitative models attempting to explain or predict animal behaviour should be able to accurately simulate behaviour in such complex, real-world conditions. Female crickets locate potential mates in choruses of simultaneously calling males. In the present study, we have tested field cricket acoustic orientation behaviour in complex acoustic conditions in the field and also successfully predicted female orientation and paths under these conditions using a simulation model based on auditory physiology. Such simulation models can provide powerful tools to predict and dissect patterns of behaviour in complex, natural environments.

Males of several acoustically communicating orthopteran species form spatially and temporally structured choruses. We investigated whether male field crickets of the species Plebeiogryllus guttiventris formed choruses in the field. Males formed spatial aggregations and showed fidelity to a calling site within a night, forming stable choruses. Within aggregations, the acoustic ranges of males overlapped considerably. We tested whether males within hearing range of each other interacted acoustically. The chirps of simultaneously calling males were aphasic with respect to each other and showed no significant alternation or synchrony of calls. Some individuals changed temporal features of their calling songs such as chirp durations and chirp rates in response to a simultaneously calling neighbour. The implications of these results for female mate choice are discussed.

The sequencing of the Mycobacterium tuberculosis (MTB) H37Rv genome has facilitated deeper insights into the biology of MTB, yet the functions of many MTB proteins are unknown. We have used sensitive profile-based search procedures to assign functional and structural domains to infer functions of gene products encoded in MTB. These domain assignments have been made using a compendium of sequence and structural domain families. Functions are predicted for 78 % of the encoded gene products. For 69 % of these, functions can be inferred by domain assignments. The functions for the rest are deduced from their homology to proteins of known function. Superfamily relationships between families of unknown and known structures have increased structural information by approximately 11%. Remote similarity detection methods have enabled domain assignments for 1325 'hypothetical proteins'. The most populated families in MTB are involved in lipid metabolism, entry and survival of the bacillus in host. Interestingly, for 353 proteins, which we refer to as MTB-specific, no homologues have been identified. Numerous, previously unannotated, hypothetical proteins have been assigned domains and some of these could perhaps be the possible chemotherapeutic targets. MTB-specific proteins might include factors responsible for virulence. Importantly, these assignments could be valuable for experimental endeavors. The detailed results are publicly available at http://hodgkin.mbu.iisc.ernet.in/~dots.

Active amplification in auditory systems is a unique and sophisticated mechanism that expends energy in amplifying the mechanical input to the auditory system, to increase its sensitivity and acuity. Although known for decades from vertebrates, active auditory amplification was only discovered in insects relatively recently. It was first discovered from two dipterans, mosquitoes and flies, who hear with their light and compliant antennae; only recently has it been observed in the stiffer and heavier tympanal ears of an orthopteran. The discovery of active amplification in two distinct insect lineages with independently evolved ears, suggests that the trait may be ancestral, and other insects may possess it as well. This opens up extensive research possibilities in the field of acoustic communication, not just in auditory biophysics, but also in behaviour and neurobiology. The scope of this review is to establish benchmarks for identifying the presence of active amplification in an auditory system and to review the evidence we currently have from different insect ears. I also review some of the models that have been posited to explain the mechanism, both from vertebrates and insects and then review the current mechanical, neurobiological and genetic evidence for each of these models.

Animal ears are exquisitely adapted to capture sound energy and perform signal analysis. Studying the ear of the locust, we show how frequency signal analysis can be performed solely by using the structural features of the tympanum. Incident sound waves generate mechanical vibrational waves that travel across the tympanum. These waves shoal in a tsunami-like fashion, resulting in energy localization that focuses vibrations onto the mechanosensory neurons in a frequency-dependent manner. Using finite element analysis, we demonstrate that two mechanical properties of the locust tympanum, distributed thickness and tension, are necessary and sufficient to generate frequency-dependent energy localization.

Despite their small size, some insects, such as crickets, can produce high amplitude mating songs by rubbing their wings together. By exploiting structural resonance for sound radiation, crickets broadcast species-specific songs at a sharply tuned frequency. Such songs enhance the range of signal transmission, contain information about the signaler’s quality, and allow mate choice. The production of pure tones requires elaborate structural mechanisms that control and sustain resonance at the species-specific frequency. Tree crickets differ sharply from this scheme. Although they use a resonant system to produce sound, tree crickets can produce high amplitude songs at different frequencies, varying by as much as an octave. Based on an investigation of the driving mechanism and the resonant system, using laser Doppler vibrometry and finite element modeling, we show that it is the distinctive geometry of the crickets’ forewings (the resonant system) that is responsible for their capacity to vary frequency. The long, enlarged wings enable the production of high amplitude songs; however, as a mechanical consequence of the high aspect ratio, the resonant structures have multiple resonant modes that are similar in frequency. The drive produced by the singing apparatus cannot, therefore, be locked to a single frequency, and different resonant modes can easily be engaged, allowing individual males to vary the carrier frequency of their songs. Such flexibility in sound production, decoupling body size and song frequency, has important implications for conventional views of mate choice, and offers inspiration for the design of miniature, multifrequency, resonant acoustic radiators.

The ears of the desert locust perform frequency analysis by using vibrational waves that travel across its heterogeneous eardrum. Similar to frequency separation achieved by the basilar membrane of the human cochlea, the locust eardrum localizes frequency components at specific anatomical eardrum locations where specific neural mechanoreceptors make physical contact with the eardrum. This is the first example of energy localization in a peripheral receptor; the mechanical basis for such behavior is discussed.

SUMMARYAnimals communicate in non-ideal and noisy conditions. The primary method they use to improve communication efficiency is sender-receiver matching: the receiver's sensory mechanism filters the impinging signal based on the expected signal. In the context of acoustic communication in crickets, such a match is made in the frequency domain. The males broadcast a mate attraction signal, the calling song, in a narrow frequency band centred on the carrier frequency (CF), and the females are most sensitive to sound close to this frequency. In tree crickets, however, the CF changes with temperature. The mechanisms used by female tree crickets to accommodate this change in CF were investigated at the behavioural and biomechanical level. At the behavioural level, female tree crickets were broadly tuned and responded equally to CFs produced within the naturally occurring range of temperatures (18 to 27°C). To allow such a broad response, however, the transduction mechanisms that convert sound into mechanical and then neural signals must also have a broad response. The tympana of the female tree crickets exhibited a frequency response that was even broader than suggested by the behaviour. Their tympana vibrate with equal amplitude to frequencies spanning nearly an order of magnitude. Such a flat frequency response is unusual in biological systems and cannot be modelled as a simple mechanical system. This feature of the tree cricket auditory system not only has interesting implications for mate choice and species isolation but may also prove exciting for bio-mimetic applications such as the design of miniature low frequency microphones.

SUMMARYField cricket females localize one of many singing males in the field in closed-loop multi-source conditions. To understand this behaviour, field cricket phonotaxis was investigated in a closed-loop walking phonotaxis paradigm, in response to two simultaneously active speakers playing aphasic calling songs. Female phonotactic paths were oriented towards the louder sound sources, but showed great inter-individual variability. Decisions made in the initial phases were correlated with the overall directions of the paths. Interestingly, the sound pressure levels of stimuli did not greatly influence several features of phonotactic paths such as sinuosity, walking bout lengths and durations. In order to ascertain the extent of our understanding of walking phonotaxis, a stochastic model was used to simulate the behaviour observed in the experiment. The model incorporated data from the experiment and our current understanding of field cricket auditory physiology. This model, based on stochastic turning towards the louder side, successfully recaptured several qualitative and quantitative features of the observed phonotactic paths. The simulation also reproduced the paths observed in a separate outdoor field experiment. Virtual crickets that were unilaterally deafened or had poor ear directionality exhibited walking paths similar to those observed in previous experiments.

Members of a superfamily of proteins could result from divergent evolution of homologues with insignificant similarity in the amino acid sequences. A superfamily relationship is detected commonly after the three-dimensional structures of the proteins are determined using X-ray analysis or NMR. The SUPFAM database described here relates two homologous protein families in a multiple sequence alignment database of either known or unknown structure. The present release (1.1), which is the first version of the SUPFAM database, has been derived by analysing Pfam, which is one of the commonly used databases of multiple sequence alignments of homologous proteins. The first step in establishing SUPFAM is to relate Pfam families with the families in PALI, which is an alignment database of homologous proteins of known structure that is derived largely from SCOP. The second step involves relating Pfam families which could not be associated reliably with a protein superfamily of known structure. The profile matching procedure, IMPALA, has been used in these steps. The first step resulted in identification of 1280 Pfam families (out of 2697, i.e. 47%) which are related, either by close homologous connection to a SCOP family or by distant relationship to a SCOP family, potentially forming new superfamily connections. Using the profiles of 1417 Pfam families with apparently no structural information, an all-against-all comparison involving a sequence-profile match using IMPALA resulted in clustering of 67 homologous protein families of Pfam into 28 potential new superfamilies. Expansion of groups of related proteins of yet unknown structural information, as proposed in SUPFAM, should help in identifying 'priority proteins' for structure determination in structural genomics initiatives to expand the coverage of structural information in the protein sequence space. For example, we could assign 858 distinct Pfam domains in 2203 of the gene products in the genome of Mycobacterium tubercolosis. Fifty-one of these Pfam families of unknown structure could be clustered into 17 potentially new superfamilies forming good targets for structural genomics. SUPFAM database can be accessed at http://pauling.mbu.iisc.ernet.in/similar tosupfam.

The sequencing of the Mycobacterium tuberculosis (MTB) H37Rv genome has facilitated deeper insights into the biol. of MTB, yet the functions of many MTB proteins are unknown. We have used sensitive profile-based search procedures to assign functional and structural domains to infer functions of gene products encoded in MTB. These domain assignments have been made using a compendium of sequence and structural domain families. Functions are predicted for 78% of the encoded gene products. For 69% of these, functions can be inferred by domain assignments. The functions for the rest are deduced from their homology to proteins of known function. Superfamily relationships between families of unknown and known structures have increased structural information by \sim 11%. Remote similarity detection methods have enabled domain assignments for 1325 'hypothetical proteins'. The most populated families in MTB are involved in lipid metabolism entry and survival of the bacillus in host. Interestingly, for 353 proteins, which we refer to as MTB-specific, no homologs have been identified. Numerous, previously unannotated, hypothetical proteins have been assigned domains and some of these could perhaps be the possible chemotherapeutic targets. MTB-specific proteins might include factors responsible for virulence. Importantly, these assignments could be valuable for experimental endeavors. The detailed results are publicly available at http://hodgkin.mbu.iisc.ernet.in/ \sim dots