Distribution of sound pressure around a singing cricket: radiation pattern and asymmetry in the sound field

Male field crickets generate calls to attract distant females through tegminal stridulation: the rubbing together of the overlying right wing which bears a file of cuticular teeth against the underlying left wing which carries a sclerotised scraper. During stridulation, specialised areas of membrane on both wings are set into oscillating vibrations to produce acoustic radiation. The location of females is unknown to the calling males and thus increasing effective signal range in all directions will maximise transmission effectiveness. However, producing an omnidirectional sound field of high sound pressure levels may be problematic due to the mechanical asymmetry found in this sound generation system. Mechanical asymmetry occurs by the right wing coming to partially cover the left wing during the closing stroke phase of stridulation. As such, it is hypothesised that the sound field on the left-wing side of the animal will contain lower sound pressure components than on the right-wing side as a result of this coverage. This hypothesis was tested using a novel method to accurately record a high resolution, three dimensional mapping of sound pressure levels around restrained Gryllus bimaculatus field crickets singing under pharmacological stimulation. The results indicate that a bilateral asymmetry is present across individuals, with greater amplitude components present in the right wing side of the animal. Individual variation in sound pressure to either the right or left-wing side is also observed. However, statistically significant differences in bilateral sound field asymmetry as presented here may not affect signalling in the field

Mechanisms of high-frequency song generation in brachypterous crickets and the role of ghost frequencies

Sound production in crickets relies on stridulation, the well-understood rubbing together of a pair of specialised wings. As the file of one wing slides over the scraper of the other, a series of rhythmic impacts cause harmonic oscillations, usually resulting in the radiation of pure tones delivered at low frequencies (2-8 kHz). In the short winged crickets of the Lebinthini tribe, acoustic communication relies on signals with remarkably high frequencies (> 8 kHz) and rich harmonic content. Using several species of the subfamily Eneopterinae, we characterise the morphological and mechanical specialisations supporting the production of high frequencies, and demonstrate that higher harmonics are exploited as dominant frequencies. These specialisations affect the structure of the stridulatory file, the motor control of stridulation and the resonance of the sound radiator. We place these specialisations in a phylogenetic framework and show that they serve to exploit high frequency vibrational modes pre-existing in the phylogenetic ancestor. In Eneopterinae, the lower frequency components are harmonically related to the dominant peak, suggesting they are relicts of ancestral carrier frequencies. Yet, such ghost frequencies still occur in the wings' free resonances, highlighting the fundamental mechanical constraints of sound radiation. These results support the hypothesis that such high frequency songs evolved stepwise, by a form of punctuated evolution which could be related to functional constraints, rather than by the progressive increase of the ancestral fundamental frequency

Digging the optimum pit: Antlions, spirals and spontaneous stratification

Most animal traps are constructed from self-secreted silk, so antlions are rare among trap builders because they use only materials found in the environment. We show how antlions exploit the properties of the substrate to produce very effective structures in the minimum amount of time. Our modelling demonstrates how antlions: (i) exploit self-stratification in granular media differentially to expose deleterious large grains at the bottom of the construction trench where they can be ejected preferentially, and (ii) minimize completion time by spiral rather than central digging. Both phenomena are confirmed by our experiments. Spiral digging saves time because it enables the antlion to eject material initially from the periphery of the pit where it is less likely to topple back into the centre. As a result, antlions can produce their pits—lined almost exclusively with small slippery grains to maximize powerful avalanches and hence prey capture—much more quickly than if they simply dig at the pit’s centre. Our demonstration, for the first time to our knowledge, of an animal using self-stratification in granular media exemplifies the sophistication of extended phenotypes even if they are only formed from material found in the animal’s environment

Functional morphology of tegmina-based stridulation in the relict species Cyphoderris monstrosa (Orthoptera: Ensifera: Prophalangopsidae)

Male grigs, bush-crickets and field crickets produce mating calls by tegminal stridulation: the scraping together of modified forewings functioning as sound generators. Bush- (Tettigoniidae) and field-crickets (Gryllinae) diverged some 240 million years ago, with each lineage developing unique characteristics in wing morphology and the associated mechanics of stridulation. The grigs (Prophalangopsidae), a relict lineage more closely related to bush crickets than to field-crickets, are believed to retain plesiomorphic features of wing morphology. The wing cells widely involved in sound production, such as the harp and mirror, are comparatively small, poorly delimited and/or partially filled with cross-veins. Such morphology is similarly observed in the earliest stridulating ensiferans, for which stridulatory mechanics remains poorly understood. The grigs, therefore, are of major importance to investigate the early evolutionary stages of tegminal stridulation, a critical innovation in the evolution of the Orthoptera. The aim of this study is to appreciate the degree of specialisation on grig forewings, through identification of sound radiating area areas and their properties. For well-grounded comparisons, homologies in wing venation (and associated areas) of grigs and bush-crickets are re-evaluated. Then, using direct evidence, this study confirms the mirror cell, in association with two other areas (termed ‘neck’ and ‘pre-mirror’), as the acoustic resonator in the grig Cyphoderris monstrosa. Despite the use of largely symmetrical resonators, as found in field-crickets, analogous features of stridulatory mechanics are observed between C. monstrosa and bush-crickets. Both morphology and function in grigs represents transitional stages between unspecialised forewings and derived conditions observed in modern species

Jumping without Using Legs: The Jump of the Click-Beetles (Elateridae) Is Morphologically Constrained

To return to their feet, inverted click-beetles (Elateridae) jump without using their legs. When a beetle is resting on its dorsal side, a hinge mechanism is locked to store elastic energy in the body and releases it abruptly to launch the beetle into the air. While the functional morphology of the jumping mechanism is well known, the level of control that the beetle has over this jumping technique and the mechanical constraints governing the jumps are not entirely clear. Here we show that while body rotations in air are highly variable, the jumps are morphologically constrained to a constant “takeoff” angle (79.9°±1.56°, n = 9 beetles) that directs 98% of the jumping force vertically against gravity. A physical-mathematical model of the jumping action, combined with measurements from live beetle, imply that the beetle may control the speed at takeoff but not the jumping angle. In addition, the model shows that very subtle changes in the exact point of contact with the ground can explain the vigorous rotations of the body seen while the beetle is airborne. These findings suggest that the evolution of this unique non-legged jumping mechanism resulted in a jumping technique that is capable of launching the body high into the air but it is too constrained and unstable to allow control of body orientation at landing

The Mechanism of Tuning of The Mole Cricket Singing Burrow

1. Experimental and theoretical studies on the acoustics of the singing burrow of the mole cricket Gryllotalpa australis are reported. 2. The burrow typically consists of a bulb about 26 mm long and 20 mm in diameter, connected through a constriction of diameter about 10 mm to a flaring horn with length about 40 mm and equivalent mouth diameter about 34 mm. The mouth geometry of the burrow differs from one species to another, and the aperture may be either single, double or even multiple. The end of the bulb opposite the horn connects to a narrow exit tunnei of diameter about 8 mm and length up to 1 m. The singing cricket positions itself close to the constriction between the bulb and the horn and produces a song with a frequency around 2.5 kHz. 3. Measurements of sound pressure within the burrow when it is excited by an external sound source at the song frequency show a pressure minimum at the constriction and an amplitude and phase distribution that is consistent with resonance of the burrow at its second modal frequency. The burrow is approximately three-quarters of a wavelength long at this frequency. The same result is obtained when the burrow is excited by a dipole source located near the constriction. 4. Non-parametric model calculations confirm this conclusion and also give broad agreement with the measured response curves over a frequency range from about 1.5 to 5 kHz. The calculated curves indicate an additional resonance at about 1.2 kHz associated with the first mode of the burrow—the Helmholtz or Klipsch resonance—which is apparently not utilised by the insect. This detail is consistent with earlier measurements, and is also supported by measured responses reported here that show an increase in sound pressure with decreasing frequency below 2 kHz as predicted by the model. 5. The measured performance of the burrow is broadly consistent with the model. According to the model, the burrow geometry is close to optimal for maximal sound power radiation

The rhizoferrin biosynthetic gene in the fungal pathogen Rhizopus delemar is a novel member of the NIS gene family

This work was supported by the Natural Sciences and Engineering Research Council of Canada award to MM (grant number 611181). C. Carroll thanks Simon Fraser University for a travel and research award.Iron is essential for growth and in low iron environments such as serum many bacteria and fungi secrete ferric iron-chelating molecules called siderophores. All fungi produce hydroxamate siderophores with the exception of Mucorales fungi, which secrete rhizoferrin, a polycarboxylate siderophore. Here we investigated the biosynthesis of rhizoferrin by the opportunistic human pathogen, Rhizopus delemar. We searched the genome of R. delemar 99–880 for a homologue of the bacterial NRPS-independent siderophore (NIS) protein, SfnaD that is involved in biosynthesis of staphyloferrin A in Staphylococcus aureus. A protein was identified in R. delemar with 22% identity and 37% similarity with SfnaD, containing an N-terminal IucA/IucC family domain, and a C-terminal conserved ferric iron reductase FhuF-like transporter domain. Expression of the putative fungal rhizoferrin synthetase (rfs) gene was repressed by iron. The rfs gene was cloned and expressed in E.coli and siderophore biosynthesis from citrate and diaminobutane was confirmed using high resolution LC–MS. Substrate specificity was investigated showing that Rfs produced AMP when oxaloacetic acid, tricarballylic acid, ornithine, hydroxylamine, diaminopentane and diaminopropane were employed as substrates. Based on the production of AMP and the presence of a mono-substituted rhizoferrin, we suggest that Rfs is a member of the superfamily of adenylating enzymes. We used site-directed mutagenesis to mutate selected conserved residues predicted to be in the Rfs active site. These studies revealed that H484 is essential for Rfs activity and L544 may play a role in amine recognition by the enzyme. This study on Rfs is the first characterization of a fungal NIS enzyme. Future work will determine if rhizoferrin biosynthesis is required for virulence in Mucorales fungi.PostprintPeer reviewe

YYY Dynamic Tuning of Viscoelastic Hydrogels with Carbonyl Iron Microparticles Reveals the Rapid Response of Cells to Three-Dimensional Substrate Mechanics

Current methods to dynamically tune three-dimensional hydrogel mechanics require specific chemistries and substrates that make modest, slow, and often irreversible changes in their mechanical properties, exclude the use of protein-based scaffolds, or alter the hydrogel microstructure and pore size. Here, we rapidly and reversibly alter the mechanical properties of hydrogels consisting of extracellular matrix proteins and proteoglycans by adding carbonyl iron microparticles (MPs) and applying external magnetic fields. This approach drastically alters hydrogel mechanics: rheology reveals that application of a 4000 Oe magnetic field to a 5 mg/mL collagen hydrogel containing 10 wt % MPs increases the storage modulus from approximately 1.5 to 30 kPa. Cell morphology experiments show that cells embedded within these hydrogels rapidly sense the magnetically induced changes in ECM stiffness. Ca2+ transients are altered within seconds of stiffening or subsequent softening, and slower but still dynamic changes occur in YAP nuclear translocation in response to time-dependent application of a magnetic field. The near instantaneous change in hydrogel mechanics provides new insight into the effect of changing extracellular stiffness on both acute and chronic changes in diverse cell types embedded in protein-based scaffolds. Due to its flexibility, this method is broadly applicable to future studies interrogating cell mechanotransduction in three-dimensional substrates