Electroreception in marine fishes: chondrichthyans

Electroreception in marine fishes occurs across a variety of taxa and is best understood in the chondrichthyans (sharks, skates, rays, and chimaeras). Here, we present an up-to-date review of what is known about the biology of passive electroreception and we consider how electroreceptive fishes might respond to electric and magnetic stimuli in a changing marine environment. We briefly describe the history and discovery of electroreception in marine Chondrichthyes, the current understanding of the passive mode, the morphological adaptations of receptors across phylogeny and habitat, the physiological function of the peripheral and central nervous system components, and the behaviours mediated by electroreception. Additionally, whole genome sequencing, genetic screening and molecular studies promise to yield new insights into the evolution, distribution, and function of electroreceptors across different environments. This review complements that of electroreception in freshwater fishes in this special issue, which provides a comprehensive state of knowledge regarding the evolution of electroreception. We conclude that despite our improved understanding of passive electroreception, several outstanding gaps remain which limits our full comprehension of this sensory modality. Of particular concern is how electroreceptive fishes will respond and adapt to a marine environment that is being increasingly altered by anthropogenic electric and magnetic fields.     

Prey detection mechanism of elasmobranchs

Elasmobranchs can detect a little amount of electric fields and they have characteristic approach strategies to find an electric dipole source generated by prey or conspecifics. They appear to align the body at a constant angle with the current flow line of the electric field while swimming towards prey. However, it has not been studied how they process the perception of electric fields for the approach behaviour or what kind of neural mechanism is used. We use a model of electrosensory perception with electrodynamics and explore a possible approach mechanism based on the sensory landscape distributed on electroreceptors. This paper presents that elasmobranchs can estimate the direction of the electric field by swaying their head, which will be a basis information for their particular approach behaviour. A velocity profile of voltage gradients and intensity difference among the ampullary clusters can be another cues to detect a prey source.     

A video and photographic study of aggregation,swimming and respiratory behaviour changes in the Grey Nurse Shark (Carcharias taurus) in response to the presence of SCUBA divers

The Grey Nurse Shark (Carcharias taurus) is a popular attraction for shark eco-tourism using SCUBA. The species is also ‘globally Vulnerable’ (IUCN 2008. List of Threatened Species. www.iucnredlist.org/). Magic Point (off Maroubra) in Sydney is favoured by recreational SCUBA divers wishing to observe these sharks. The objective of this study was to experimentally test the level of the activities of recreational SCUBA divers on shark behaviour. This study assessed the shark responses to diver group size (4, 8 and 12), time of day (am, noon and pm) and diver distance from the sharks (3?m and 6?m). The study found that diver activity does affect the aggregation, swimming and respiratory behaviour of sharks at this site, albeit at short-term levels. Diver group size had no significant effect on shark aggregation, but the proximity of divers to the sharks was crucial. Shark distribution in the cave changed significantly in the presence of divers at 3?m distance from the cave, but stayed unchanged at 6?m. This was particularly apparent in the presence of large groups of 12 divers at 3?m distance when sharks increased their swim speed and ventilation mechanism from ‘active’ to ‘RAM’ ventilation. Such change coincided with a sudden decrease in ventilation frequency. Our research suggests that these effects are short-term and that sharks resume their behaviour once the divers retreat. If divers abide by the current code of practice for diving at this site, it is unlikely that their activities will substantially impact Grey Nurse Sharks in the long term.     

Ensemble modeling of the potential distribution of the whale shark in the Atlantic Ocean

The whale shark (Rhincodon typus) is an endangered marine fish species which can be adversely affected by the fishing activities of the industrial purse seine fleet targeting tropical tuna. Tuna tend to aggregate around all types of floating objects, including whale sharks. We analyzed and modeled the spatial distribution and environmental preferences of whale sharks based on the presence and absence data from fishing observations in the Atlantic Ocean. We used a thorough multialgorithm analysis, based on a new presence–absence dataset, and endeavored to follow the most recent recommendations on best practices in species distribution modeling. First, we selected a subset of relevant variables using a generalized linear model that addressed multicollinearity, statistical errors, and information criteria. We then used the selected variables to build a model ensemble including 19 different algorithms. After eliminating models with insufficient performance, we assessed the potential distribution of whale sharks using the mean of the predictions of the selected models. We also assessed the variance among the predictions of different algorithms, in order to identify areas with the highest model consensus. The results show that several coastal regions and warm shallow currents, such as the Gulf Stream and the Canary and Benguela currents, are the most suitable areas for whale sharks under current environmental conditions. Future environmental projections for the Atlantic Ocean suggest that some of the suitable regions will shift northward, but current concentration areas will continue to be suitable for whale shark, although with less productivity, which could have negative consequences for conservation of the species. We discuss the implications of these predictions for the conservation and management of this charismatic marine species.     

Modeling and measuring lateral line excitation patterns to changing dipole source locations

In order to determine excitation patterns to the lateral line system from a nearby 50 Hz oscillating sphere, dipole flow field equations were used to model the spatial distribution of pressures along a linear array of lateral line canal pores. Modeled predictions were then compared to pressure distributions measured for the same dipole source with a miniature hydrophone placed in a small test tank used for neurophysiological experiments. Finally, neural responses from posterior lateral line nerve fibers in the goldfish were measured in the test tank to demonstrate that modeled and measured pressure gradient patterns were encoded by the lateral line periphery. Response patterns to a 50 Hz dipole source that slowly changed location along the length of the fish included (1) peaks and valleys in spike-rate responses corresponding to changes in pressure gradient amplitudes, (2) 180° phase-shifts corresponding to reversals in the direction of the pressure gradient and (3) distance-dependent changes in the locations of peaks, valleys and 180° phase-shifts. Modeled pressure gradient patterns also predict that the number of neural amplitude peaks and phase transitions will vary as a function of neuromast orientation and axis of source oscillation. The faithful way in which the lateral line periphery encodes pressure gradient patterns has implications for how source location and distance might be encoded by excitation patterns in the CNS. Phase-shift information may be important for (1) inhibitory/excitatory sculpting of receptive fields and (2) unambiguously encoding source distance so that increases in source distance are not confused with decreases in source amplitude.     

The effect of translocating cylindrical particles on the ionic current through a nanopore

The electric field-induced translocation of cylindrical particles through nanopores with circular cross sections is studied theoretically. The coupled Nernst-Planck equations (multi-ion model, MIM) for the concentration fields of the ions in solution and the Stokes equation for the flow field are solved simultaneously. The predictions of the multi-ion model are compared with the predictions of two simplified models based on the Poisson-Boltzmann equation (PBM) and the Smoluchowski's slip velocity (SVM). The concentration field, the ionic current though the pore, and the particle's velocity are computed as functions of the particle's size, location, and electric charge; the pore's size and electric charge; the electric field intensity; and the bulk solution's concentration. In qualitative agreement with experimental data, the MIM predicts that, depending on the bulk solution's concentration, the translocating particle may either block or enhance the ionic current. When the thickness of the electric double layer is relatively large, the PBM and SVM predictions do not agree with the MIM predictions. The limitations of the PBM and SVM are delineated. The theoretical predictions are compared with and used to explain experimental data pertaining to the translocation of DNA molecules through nanopores.     

To go or not to go with the flow: Environmental influences on whale shark movement patterns

Seven whale sharks were tracked using satellite-linked tags from Ningaloo Reef, off northern Western Australia, following tagging in April and June 2002 and April-May 2005. We investigated how the movements of those whale shark tracks were influenced by geostrophic surface currents during sequential one-week periods by using a passive diffusion model parameterised with observed starting locations of the sharks and weekly maps of surface current velocity and direction (derived from altimetry). We compared the outputs from the passive diffusion model and maps of chlorophyll-a concentration (SeaWiFs/MODIS) and with the actual tracks of the sharks using GIS and generalized linear mixed-effects models (GLMM). The GLMM indicated very little support for passive diffusion with sea-surface ocean currents influencing whale shark distributions in the north eastern Indian Ocean. Moreover, the sharks' movements correlated only weakly with the spatial distribution of sea-surface chlorophyll-a concentrations. The seven whale sharks had average swimming speeds comparable with those recorded in other satellite tracking studies of this species. Swimming speeds of the seven sharks were similar to those reported in previous studies and up to three times greater than the maximum sea-surface current velocities that the sharks encountered while traversing into lower southerly latitudes (moving northward towards the equator). Our results indicate that whale sharks departing from Ningaloo travel actively and independently of near-surface currents where they spend most of their time despite additional metabolic costs of this behaviour.     

Empirical and theoretical dosimetry in support of whole body radio frequency (RF) exposure in seated human volunteers at 220 MHz

This study reports the dosimetry performed to support an experiment that measured physiological responses of seated volunteer human subjects exposed to 220 MHz fields. Exposures were performed in an anechoic chamber which was designed to provide uniform fields for frequencies of 100 MHz or greater. A vertical half-wave dipole with a 90 degrees reflector was used to optimize the field at the subject's location. The vertically polarized E field was incident on the dorsal side of the phantoms and human volunteers. The dosimetry plan required measurement of stationary probe drift, field strengths as a function of distance, electric and magnetic field maps at 200, 225, and 250 cm from the dipole antenna, and specific absorption rate (SAR) measurements using a human phantom, as well as theoretical predictions of SAR with the finite difference time domain (FDTD) method. A NBS (National Bureau of Standards, now NIST, National Institute of Standards and Technology, Boulder, CO) 10 cm loop antenna was positioned 150 cm to the right, 100 cm above and 60 cm behind the subject (toward the transmitting antenna) and was read prior to each subject's exposure and at 5 min intervals during all RF exposures. Transmitter stability was determined by measuring plate voltage, plate current, screen voltage and grid voltage for the driver and final amplifiers before and at 5 min intervals throughout the RF exposures. These dosimetry measurements assured accurate and consistent exposures. FDTD calculations were used to determine SAR distribution in a seated human subject. This study reports the necessary dosimetry to precisely control exposure levels for studies of the physiological consequences of human volunteer exposures to 220 MHz.     

Ion acceleration in a dipole vortex in a laser plasma corona

Particle-in-cell simulations show that the inhomogeneity scale of the plasma produced in the interaction of high-power laser radiation with gas targets is of fundamental importance for ion acceleration. In a plasma slab with sharp boundaries, the quasistatic magnetic field and the associated electron vortex structure produced by fast electron beams both expand along the slab boundary in a direction perpendicular to the plasma density gradient, forming an extended region with a quasistatic electric field, in which the ions are accelerated. In a plasma with a smooth density distribution, the dipole magnetic field can propagate toward the lower plasma density in the propagation direction of the laser pulse. In this case, the electron density in an electric current filament at the axis of the magnetic dipole decreases to values at which the charge quasineutrality condition fails to hold. In electric fields generated by this process, the ions are accelerated to energies substantially higher than those characteristic of plasma configurations with sharp boundaries.     

Probing the effect of electric field in 9,10-dimethoxy-2,6-bis(2-p-tolylethynyl)anthracene molecular nanowire using quantum chemical and charge density analysis

The effect of electric field (EF) in a newly designed molecular nanowire 9,10-dimethoxy-2,6-bis(2-p-tolylethynyl)anthracene has been analysed theoretically from the structural and electronic charge transport properties using quantum chemical and charge density calculations. The applied EF (0–0.36 VÅ^? 1) alters the molecular conformation, charge density distribution, electrostatic properties and the electronic energy levels of the molecule. Furthermore, the applied EF decreases the highest occupied molecular orbital–lowest unoccupied molecular orbital gap significantly from 1.775 to 0.258 eV and it also induces polarisation in the molecule, which leads to increase the dipole moment of the molecule. The electrostatic potential for various levels of applied EF reveals the charge-accumulated regions of the molecule. The I–V characteristics of the molecule have been studied against various applied fields using Landauer formalism.     

Visualizing ion relaxation in the transport of short DNA fragments

Ion relaxation plays an important role in a wide range of phenomena involving the transport of charged biomolecules. Ion relaxation is responsible for reducing sedimentation and diffusion constants, reducing electrophoretic mobilities, increasing intrinsic viscosities, and, for biomolecules that lack a permanent electric dipole moment, provides a mechanism for orienting them in an external electric field. Recently, a numerical boundary element method was developed to solve the coupled Navier-Stokes, Poisson, and ion transport equations for a polyion modeled as a rigid body of arbitrary size, shape, and charge distribution. This method has subsequently been used to compute the electrophoretic mobilities and intrinsic viscosities of a number of model proteins and DNA fragments. The primary purpose of the present work is to examine the effect of ion relaxation on the ion density and fluid velocity fields around short DNA fragments (20 and 40 bp). Contour density as well as vector field diagrams of the various scalar and vector fields are presented and discussed at monovalent salt concentrations of 0.03 and 0.11 M. In addition, the net charge current fluxes in the vicinity of the DNA fragments at low and high salt concentrations are briefly examined and discussed.     

A model of the electric field of the brain at EEG and microwave frequencies

A simple calculation of the current dipole moment of the extracellular electric field of the cortex is proposed; it is based on the dipole layer model. The model is extended to the range of microwave frequencies. Arguments in favor of emission of microwave radiation by the dendritic membranes of pyramidal neurons are presented and the strength of the radiative electric field at a distance from the head is calculated and discussed.     

Exaggerated trans-membrane charge of ammonium transporters in nutrient-poor marine environments

Transporter proteins are a vital interface between cells and their environment. In nutrient-limited environments, microbes with transporters that are effective at bringing substrates into their cells will gain a competitive advantage over variants with reduced transport function. Microbial ammonium transporters (Amt) bring ammonium into the cytoplasm from the surrounding periplasm space, but diagnosing Amt adaptations to low nutrient environments solely from sequence data has been elusive. Here, we report altered Amt sequence amino acid distribution from deep marine samples compared to variants sampled from shallow water in two important microbial lineages of the marine water column community—Marine Group I Archaea (Thermoproteota) and the uncultivated gammaproteobacterial lineage SAR86. This pattern indicates an evolutionary pressure towards an increasing dipole in Amt for these clades in deep ocean environments and is predicted to generate stronger electric fields facilitating ammonium acquisition. This pattern of increasing dipole charge with depth was not observed in lineages capable of accessing alternative nitrogen sources, including the abundant alphaproteobacterial clade SAR11. We speculate that competition for ammonium in the deep ocean drives transporter sequence evolution. The low concentration of ammonium in the deep ocean is therefore likely due to rapid uptake by Amts concurrent with decreasing nutrient flux.     

Empirical and theoretical dosimetry in support of whole body resonant RF exposure (100 MHz) in human volunteers

This study reports the dosimetry performed to support an experiment that measured physiological responses of volunteer human subjects exposed to the resonant frequency for a seated human adult at 100 MHz. Exposures were performed in an anechoic chamber which was designed to provide uniform fields for frequencies of 100 MHz or greater. A half wave dipole with a 90 degrees reflector was used to optimize the field at the subject location. The dosimetry plan required measurement of transmitter harmonics, stationary probe drift, field strengths as a function of distance, electric and magnetic field maps at 200, 225, and 250 cm from the dipole antenna, and specific absorption rate (SAR) measurements using a human phantom, as well as theoretical predictions of SAR with the finite difference time domain (FDTD) method. On each exposure test day, a measurement was taken at 225 cm on the beam centerline with a NBS E field probe to assure consistently precise exposures. A NBS 10 cm loop antenna was positioned 150 cm to the right, 100 cm above, and 60 cm behind the subject and was read at 5 min intervals during all RF exposures. These dosimetry measurements assured accurate and consistent exposures. FDTD calculations were used to determine SAR distribution in a seated human subject. This study reports the necessary dosimetry for work on physiological consequences of human volunteer exposures to 100 MHz.     

The physical nature of life.

Life evolved from the primeval world of physics. Sensory systems inform animals of the natural environment, enabling them to conduct responsively. The discovery of weak, DC bioelectric fields in the vicinity of aquatic organisms and the role they play in guiding sharks and rays to their prey have led to the recognition of fundamental, hitherto less well known, physical aspects of sensory biology. The inferred cybernetic algorithm of electric-field orientation in sharks and rays is highly effective and extremely robust. In orienting to the weak DC electric fields of ocean currents and to the earth's magnetic field, sharks and rays unwittingly practise the motional-electric principles that Einstein had in mind when he introduced the special theory of relativity. At the sense-organ, receptor-membrane, and ion-channel levels, the elasmobranch ampullae of Lorenzini operate on the basis of graded positive feedback driven by negative conductance, supposedly employing voltage-sensitive ion channels as the active, excitable elements. The electric sense of sharks and rays presents an exquisite implementation of the very biophysical principles that also govern the graded, much richer than all-or-none, integrative brain processes of animal and man.     

Electrosensitive spatial vectors in elasmobranch fishes: implications for source localization

The electrosense of sharks and rays is used to detect weak dipole-like bioelectric fields of prey, mates and predators, and several models propose a use for the detection of streaming ocean currents and swimming-induced fields for geomagnetic orientation. We assessed pore distributions, canal vectors, complementarity and possible evolutionary divergent functions for ampullary clusters in two sharks, the scalloped hammerhead (Sphyrna lewini) and the sandbar shark (Carcharhinus plumbeus), and the brown stingray (Dasyatis lata). Canal projections were determined from measured coordinates of each electrosensory pore and corresponding ampulla relative to the body axis. These species share three ampullary groups: the buccal (BUC), mandibular (MAN) and superficial ophthalmic (SO), which is subdivided into anterior (SOa) and posterior (SOp) in sharks. The stingray also has a hyoid (HYO) cluster. The SOp in both sharks contains the longest (most sensitive) canals with main projections in the posterior-lateral quadrants of the horizontal plane. In contrast, stingray SO canals are few and short with the posterior-lateral projections subsumed by the HYO. There was strong projection coincidence by BUC and SOp canals in the posterior lateral quadrant of the hammerhead shark, and laterally among the stingray BUC and HYO. The shark SOa and stingray SO and BUC contain short canals located anterior to the mouth for detection of prey at close distance. The MAN canals of all species project in anterior or posterior directions behind the mouth and likely coordinate prey capture. Vertical elevation was greatest in the BUC of the sandbar shark, restricted by the hammerhead cephalofoil and extremely limited in the dorsoventrally flattened stingray. These results are consistent with the functional subunit hypothesis that predicts specialized ampullary functions for processing of weak dipole and geomagnetic induced fields, and provides an anatomical basis for future experiments on central processing of different forms of relevant electric stimuli.     

Procedure using voltage-sensitive spin-labels to monitor dipole potential changes in phospholipid vesicles: The estimation of phloretin-induced conductance changes in vesicles

Summary Voltage-sensitive membrane potential probes were used to monitor currents resulting from positive or negative charge movement across small and large unilamellar phosphatidylcholine (PC) vesicles. Positive currents were measured for the paramagnetic phosphonium ion or for K⁺-valinomycin. Negative currents were indirectly measured for the anionic proton carriers CCCP and DNP by monitoring transmembrane proton currents. Phloretin, a compound that is believed to decrease dipole fields in planar bilayers, increases positive currents and decreases negative currents when added to egg PC vesicles. In these vesicles, positive currents are increased by phloretin addition to a much larger degree than CCCP currents are reduced. This asymmetry, with respect to the sign of the charge carrier, is apparently not the result of changes in the membrane dielectric constant. It is most easily explained by deeper binding minima at the membrane-solution interface for the CCCP anion, when compared to the phosphonium. The measured asymmetry and the magnitudes of the current changes are consistent with the predictions of a point dipole model. The use of potential-sensitive probes to estimate positive and negative currents, provides a methodology to monitor changes in the membrane dipole potential in vesicle systems.     

Brain stimulation using electromagnetic sources: theoretical aspects.

We prove that, at the frequencies generally proposed for extracranial stimulation of the brain, it is not possible, using any superposition of external current sources, to produce a three-dimensional local maximum of the electric field strength inside the brain. The maximum always occurs on a boundary where the conductivity jumps in value. Nevertheless, it may be possible to achieve greater two-dimensional focusing and shaping of the electric field than is currently available. Towards this goal we have used the reciprocity theorem to present a uniform treatment of the electric field inside a conducting medium produced by a variety of sources: an external magnetic dipole (current loop), an external electric dipole (linear antenna), and surface and depth electrodes. This formulation makes use of the lead fields from magneto- and electroencephalography. For the special case of a system with spherically symmetric conductivity, we derive a simple analytic formula for the electric field due to an external magnetic dipole. This formula is independent of the conductivity profile and therefore embraces spherical models with any number of shells. This explains the "insensitivity" to the skull's conductivity that has been described in numerical studies. We also present analytic formulas for the electric field due to an electric dipole, and also surface and depth electrodes, for the case of a sphere of constant conductivity.