Tracking apex marine predator movements in a dynamic ocean.

Pelagic marine predators face unprecedented challenges and uncertain futures. Overexploitation and climate variability impact the abundance and distribution of top predators in ocean ecosystems. Improved understanding of ecological patterns, evolutionary constraints and ecosystem function is critical for preventing extinctions, loss of biodiversity and disruption of ecosystem services. Recent advances in electronic tagging techniques have provided the capacity to observe the movements and long-distance migrations of animals in relation to ocean processes across a range of ecological scales. Tagging of Pacific Predators, a field programme of the Census of Marine Life, deployed 4,306 tags on 23 species in the North Pacific Ocean, resulting in a tracking data set of unprecedented scale and species diversity that covers 265,386 tracking days from 2000 to 2009. Here we report migration pathways, link ocean features to multispecies hotspots and illustrate niche partitioning within and among congener guilds. Our results indicate that the California Current large marine ecosystem and the North Pacific transition zone attract and retain a diverse assemblage of marine vertebrates. Within the California Current large marine ecosystem, several predator guilds seasonally undertake north-south migrations that may be driven by oceanic processes, species-specific thermal tolerances and shifts in prey distributions. We identify critical habitats across multinational boundaries and show that top predators exploit their environment in predictable ways, providing the foundation for spatial management of large marine ecosystems.

A novel application of multi-event modeling to estimate class segregation in a highly migratory oceanic vertebrate.

Spatial segregation of animals by class (i.e., maturity or sex) within a population due to differential rates of temporary emigration (TE) from study sites can be an important life history feature to consider in population assessment and management. However, such rates are poorly known; new quantitative approaches to address these knowledge gaps are needed. We present a novel application of multi-event models that takes advantage of two sources of detections to differentiate temporary emigration from apparent absence to quantify class segregation within a study population of double-marked (photo-identified and tagged with coded acoustic transmitters) white sharks (Carcharodon carcharias) in central California. We use this model to test if sex-specific patterns in TE result in disparate apparent capture probabilities (po ) between male and female white sharks, which can affect the observed sex ratio. The best-supported model showed a contrasting pattern of Pr(TE) from coastal aggregation sites between sexes (for males Pr[TE] = 0.015 [95% CI = 0.00, 0.31] and Pr[TE]= 0.57 [0.40, 0.72] for females), but not maturity classes. Additionally, by accounting for Pr(TE) and imperfect detection, we were able to estimate class-specific values of true capture probability (p* ) for tagged and untagged sharks. The best-supported model identified differences between maturity classes but no difference between sexes or tagging impacts (tagged mature sharks p*  = 0.55 (0.46-0.63) and sub-adult sharks p*  = 0.36 (0.25, 0.50); and untagged mature sharks p*  = 0.50 (0.39-0.61) and sub-adults p*  = 0.18 (0.10, 0.31). Estimated sex-based differences in po were linked to sex-specific differences in Pr(TE) but not in p* ; once the Pr(TE) is accounted for, the p* between sexes was not different. These results indicate that the observed sex ratio is not a consequence of unequal detectability and sex-specific values of Pr(TE) are important drivers of the observed male-dominated sex ratio. Our modeling approach reveals complex class-specific patterns in Pr(TE) and p* in a mark-recapture data set, and highlights challenges for the population modeling and conservation of white sharks in central California. The model we develop here can be used to estimate rates of temporary emigration and class segregation when two detection methods are used.

Cardiac function in an endothermic fish: cellular mechanisms for overcoming acute thermal challenges during diving.

Understanding the physiology of vertebrate thermal tolerance is critical for predicting how animals respond to climate change. Pacific bluefin tuna experience a wide range of ambient sea temperatures and occupy the largest geographical niche of all tunas. Their capacity to endure thermal challenge is due in part to enhanced expression and activity of key proteins involved in cardiac excitation-contraction coupling, which improve cardiomyocyte function and whole animal performance during temperature change. To define the cellular mechanisms that enable bluefin tuna hearts to function during acute temperature change, we investigated the performance of freshly isolated ventricular myocytes using confocal microscopy and electrophysiology. We demonstrate that acute cooling and warming (between 8 and 28°C) modulates the excitability of the cardiomyocyte by altering the action potential (AP) duration and the amplitude and kinetics of the cellular Ca(2+) transient. We then explored the interactions between temperature, adrenergic stimulation and contraction frequency, and show that when these stressors are combined in a physiologically relevant way, they alter AP characteristics to stabilize excitation-contraction coupling across an acute 20°C temperature range. This allows the tuna heart to maintain consistent contraction and relaxation cycles during acute thermal challenges. We hypothesize that this cardiac capacity plays a key role in the bluefin tunas' niche expansion across a broad thermal and geographical range.

Heart rate responses to temperature in free-swimming Pacific bluefin tuna (Thunnus orientalis).

The bluefin tuna heart remains at ambient water temperature (Ta) but must supply blood to warm regions of the body served by countercurrent vascular heat exchangers. Despite this unusual physiology, inherent difficulties have precluded an understanding of the cardiovascular responses to Ta in free-swimming bluefin tunas. We measured the heart rate (f(H)) responses of two captive Pacific bluefin tunas (Thunnus orientalis; 9.7 and 13.3 kg) over a cumulative period of 40 days. Routine f(H) during fasting in the holding tank at a Ta of 20°C was 45.1±8.0 and 40.7±6.5 beats min(-1) for Tuna 1 and Tuna 2, respectively. f(H) decreased in each fish with a Q10 temperature coefficient of 2.6 (Tuna 1) and 3.1 (Tuna 2) as Ta in the tank was slowly decreased to 15°C (~0.4°C h(-1)), despite a gradual increase in swimming speed. The same thermal challenge during digestion revealed similar thermal dependence of f(H) and indicated that the rate of visceral cooling is not buffered by the heat increment of feeding. Acutely decreasing Ta from 20 to 10°C while Tuna 1 swam in a tunnel respirometer caused a progressive increase in tail-beat frequency and oxygen consumption rate (M(O2)). f(H) of this fish decreased with a Q10 of 2.7 as Ta decreased between 20 and 15°C, while further cooling to 10°C saw a general plateau in f(H) around 35 beats min(-1) with a Q10 of 1.3. A discussion of the relationships between f(H), and haemoglobin-oxygen binding sheds further light on how bluefin cardiorespiratory systems function in a changing thermal environment.

Quantifying energy intake in Pacific bluefin tuna (Thunnus orientalis) using the heat increment of feeding.

Using implanted archival tags, we examined the effects of meal caloric value, food type (sardine or squid) and ambient temperature on the magnitude and duration of the heat increment of feeding in three captive juvenile Pacific bluefin tuna. The objective of our study was to develop a model that can be used to estimate energy intake in wild fish of similar body mass. Both the magnitude and duration of the heat increment of feeding (measured by visceral warming) showed a strong positive correlation with the caloric value of the ingested meal. Controlling for meal caloric value, the extent of visceral warming was significantly greater at lower ambient temperature. The extent of visceral warming was also significantly higher for squid meals compared with sardine meals. By using a hierarchical Bayesian model to analyze our data and treating individuals as random effects, we demonstrate how increases in visceral temperature can be used to estimate the energy intake of wild Pacific bluefin tuna of similar body mass to the individuals used in our study.

Numerical approaches to determine the interface tension of curved interfaces from free energy calculations.

A recently proposed method to obtain the surface free energy σ(R) of spherical droplets and bubbles of fluids, using a thermodynamic analysis of two-phase coexistence in finite boxes at fixed total density, is reconsidered and extended. Building on a comprehensive review of the basic thermodynamic theory, it is shown that from this analysis one can extract both the equimolar radius R(e) as well as the radius R(s) of the surface of tension. Hence the free energy barrier that needs to be overcome in nucleation events where critical droplets and bubbles are formed can be reliably estimated for the range of radii that is of physical interest. It is found that the conventional theory of nucleation, where the interface tension of planar liquid-vapor interfaces is used to predict nucleation barriers, leads to a significant overestimation, and this failure is particularly large for bubbles. Furthermore, different routes to estimate the effective radius-dependent Tolman length δ(R(s)) from simulations in the canonical ensemble are discussed. Thus we obtain an instructive exemplification of the basic quantities and relations of the thermodynamic theory of metastable droplets/bubbles using simulations. However, the simulation results for δ(R(s)) employing a truncated Lennard-Jones system suffer to some extent from unexplained finite size effects, while no such finite size effects are found in corresponding density functional calculations. The numerical results are compatible with the expectation that δ(R(s) → ∞) is slightly negative and of the order of one tenth of a Lennard-Jones diameter, but much larger systems need to be simulated to allow more precise estimates of δ(R(s) → ∞).

Warm fish with cold hearts: thermal plasticity of excitation-contraction coupling in bluefin tuna.

Bluefin tuna have a unique physiology. Elevated metabolic rates coupled with heat exchangers enable bluefin tunas to conserve heat in their locomotory muscle, viscera, eyes and brain, yet their hearts operate at ambient water temperature. This arrangement of a warm fish with a cold heart is unique among vertebrates and can result in a reduction in cardiac function in the cold despite the elevated metabolic demands of endothermic tissues. In this study, we used laser scanning confocal microscopy and electron microscopy to investigate how acute and chronic temperature change affects tuna cardiac function. We examined the temporal and spatial properties of the intracellular Ca2+ transient (Δ[Ca2+]i) in Pacific bluefin tuna (Thunnus orientalis) ventricular myocytes at the acclimation temperatures of 14°C and 24°C and at a common test temperature of 19°C. Acute (less than 5 min) warming and cooling accelerated and slowed the kinetics of Δ[Ca2+]i, indicating that temperature change limits cardiac myocyte performance. Importantly, we show that thermal acclimation offered partial compensation for these direct effects of temperature. Prolonged cold exposure (more than four weeks) increased the amplitude and kinetics of Δ[Ca2+]i by increasing intracellular Ca2+ cycling through the sarcoplasmic reticulum (SR). These functional findings are supported by electron microscopy, which revealed a greater volume fraction of ventricular SR in cold-acclimated tuna myocytes. The results indicate that SR function is crucial to the performance of the bluefin tuna heart in the cold. We suggest that SR Ca2+ cycling is the malleable unit of cellular Ca2+ flux, offering a mechanism for thermal plasticity in fish hearts. These findings have implications beyond endothermic fish and may help to delineate the key steps required to protect vertebrate cardiac function in the cold.

Temperature dependence of cardiac sarcoplasmic reticulum Ca²âº-ATPase from rainbow trout Oncorhynchus mykiss.

In this work, the temperature dependence of the sarco-endoplasmic reticulum Ca(2+) -ATPase (SERCA2) activity from rainbow trout Oncorhynchus mykiss cardiac ventricles was measured and compared with the mammalian SERCA2 isoform. The rate of ATP-dependent Ca(2+) transport catalysed by O. mykiss vesicles was totally abolished by thapsigargin and the Ca(2+) ionophore A(23187) . At warm temperatures (25 and 30° C), the SERCA2 from O. mykiss ventricles displayed the same rate of Ca(2+) uptake. At 35° C, the activity of the O. mykiss enzyme decreased after 20 min of reaction time. The rate of Ca(2+) uptake catalysed by the mammalian SERCA2 was temperature dependent exhibiting its maximal activity at 35° C. In contrast to the rate of Ca(2+) uptake, the rate of ATP hydrolysis catalysed by O. mykiss SERCA2 was not significantly different at 25 and 35° C, but the rate of ATP hydrolysis catalysed by the rat Rattus norvegicus SERCA2 isoform at 35° C was two-fold higher than at 25° C. At low temperatures (5 to 20° C), the rate of Ca(2+) uptake from O. mykiss SR was less temperature dependent than the R. norvegicus isoform, being able to sustain a high activity even at 5° C. The mean ±s.e. Q(10) values calculated from 25 to 35° C for ATP hydrolysis were 1·112 ± 0·026 (n = 3) and 2·759 ± 0·240 (n = 5) for O. mykiss and R. norvegicus, respectively. Taken together, the results show that the O. mykiss SERCA2 was not temperature dependent over the 10 to 25° C temperature interval commonly experienced by the animal in vivo. The Q(10) value of SERCA2 was significantly lower in O. mykiss than R. norvegicus which may be key for cardiac function over the wide environmental temperatures experienced in this eurythermal fish.

Shorter heels are linked with greater elastic energy storage in the Achilles tendon.

Previous research suggests that the moment arm of the m. triceps surae tendon (i.e., Achilles tendon), is positively correlated with the energetic cost of running. This relationship is derived from a model which predicts that shorter ankle moment arms place larger loads on the Achilles tendon, which should result in a greater amount of elastic energy storage and return. However, previous research has not empirically tested this assumed relationship. We test this hypothesis using an inverse dynamics approach in human subjects (n = 24) at speeds ranging from walking to sprinting. The spring function of the Achilles tendon was evaluated using specific net work, a metric of mechanical energy production versus absorption at a limb joint. We also combined kinematic and morphological data to directly estimate tendon stress and elastic energy storage. We find that moment arm length significantly determines the spring-like behavior of the Achilles tendon, as well as estimates of mass-specific tendon stress and elastic energy storage at running and sprinting speeds. Our results provide support for the relationship between short Achilles tendon moment arms and increased elastic energy storage, providing an empirical mechanical rationale for previous studies demonstrating a relationship between calcaneal length and running economy. We also demonstrate that speed and kinematics moderate tendon performance, suggesting a complex relationship between lower limb geometry and foot strike pattern.

Postprandial metabolism of Pacific bluefin tuna (Thunnus orientalis).

Specific dynamic action (SDA) is defined as the energy expended during ingestion, digestion, absorption and assimilation of a meal. This study presents the first data on the SDA response of individual tunas of any species. Juvenile Pacific bluefin tunas (Thunnus orientalis; body mass 9.7-11.0 kg; N=7) were individually fed known quantities of food consisting primarily of squid and sardine (meal energy range 1680-8749 kJ, approximately 4-13% of tuna body mass). Oxygen consumption rates (M(O2)) were measured in a swim tunnel respirometer during the postprandial period at a swimming speed of 1 body length (BL) s(-1) and a water temperature of 20 degrees C. was markedly elevated above routine levels in all fish following meal consumption [routine metabolic rate (RMR)=174+/-9 mg kg(-1) h(-1)]. The peak M(O2) during the SDA process ranged from 250 to 440 mg kg(-1) h(-1) (1.5-2.3 times RMR) and was linearly related to meal energy content. The duration of the postprandial increment in M(O2) ranged from 21 h to 33 h depending upon meal energy content. Consequently, the total energy used in SDA increased linearly with meal energy and ranged from 170 kJ to 688 kJ, such that the SDA process accounted for 9.2+/-0.7% of ingested energy across all experiments. These values suggest rapid and efficient food conversion in T. orientalis in comparison with most other fishes. Implanted archival temperature tags recorded the increment in visceral temperature (T(V)) in association with SDA. M(O2) returned to routine levels at the end of the digestive period 2-3 h earlier than T(V). The qualitative patterns in M(O2) and T(V) during digestion were similar, strengthening the possibility that archival measurements of T(V) can provide new insight into the energetics and habitat utilization of free-swimming bluefin in the natural environment. Despite efficient food conversion, SDA is likely to represent a significant component of the daily energy budget of wild bluefin tunas due to a regular and high ingestion of forage.

Multilocus phylogenetic analyses reveal that habitat selection drives the speciation of Didymozoidae (Digenea) parasitizing Pacific and Atlantic bluefin tunas.

Parasite communities of wild and reared bluefin tuna display remarkable diversity. Among these, the most prevalent and abundant are the Didymozoidae (Monticelli, 1888) (Trematoda, Digenea), considered one of the most taxonomically complex digenean families. The aim of this study was to evaluate phylogenetic structure of Didymozoidae occurring in Pacific (Thunnus orientalis) and Atlantic bluefin tuna (T. thynnus) in order to increase our knowledge of didymozoid zoogeography and identify species that could successfully be employed as biological tags for stock assessment studies. For the present analyses we used 2 nuclear ribosomal DNA loci, part of the 28S gene and the second internal transcribed spacer (ITS-2) as well as a portion of the mitochondrial cytochrome c oxidase subunit 1 gene (cox1). In most parasitic groups, morphology is the primary factor in the structuring of phylogenetic relationships. In rare examples, however, habitat has been suggested as a primary factor affecting parasite evolution. During their evolution, didymozoids have spread and inhabited a remarkable number of different sites in their hosts, colonizing exterior as well as strictly interior niches. Our data suggest that habitat selection has been the leading force in shaping didymozoid phylogenetic relationships. For 2 didymozoid species (D. wedli and D. palati), cox1 sequences indicate intraspecific differences between Mexican and Adriatic populations.

Effect of thermal acclimation on action potentials and sarcolemmal K+ channels from Pacific bluefin tuna cardiomyocytes.

To sustain cardiac muscle contractility relatively independent of temperature, some fish species are capable of temporarily altering excitation-contraction coupling processes to meet the demands of their environment. The Pacific bluefin tuna, Thunnus orientalis, is a partially endothermic fish that inhabits a wide range of thermal niches. The present study examined the effects of temperature and thermal acclimation on sarcolemmal K(+) currents and their role in action potential (AP) generation in bluefin tuna cardiomyocytes. Atrial and ventricular myocytes were enzymatically isolated from cold (14 degrees C)- and warm (24 degrees C)-acclimated bluefin tuna. APs and current-voltage relations of K(+) channels were measured using the whole cell current and voltage clamp techniques, respectively. Data were collected either at the cardiomyocytes' respective acclimation temperature of 14 or 24 degrees C or at a common test temperature of 19 degrees C (to reveal the effects of acclimation). AP duration (APD) was prolonged in cold-acclimated (CA) cardiomyocytes tested at 14 degrees C compared with 19 degrees C and in warm-acclimated (WA) cardiomyocytes tested at 19 degrees C compared with 24 degrees C. This effect was mirrored by a decrease in the density of the delayed-rectifier current (I(Kr)), whereas the density of the background inward-rectifier current (I(K1)) was unchanged. When CA and WA cardiomyocytes were tested at a common temperature of 19 degrees C, no significant effects of temperature acclimation on AP shape or duration were observed, whereas I(Kr) density was markedly increased in CA cardiomyocytes. I(K1) density was unaffected in CA ventricular myocytes but was significantly reduced in CA atrial myocytes, resulting in a depolarization of atrial resting membrane potential. Our results indicate the bluefin AP is relatively short compared with other teleosts, which may allow the bluefin heart to function at cold temperatures without the necessity for thermal compensation of APD.

The structure of the membrane systems in a novel muscle cell modified for heat production.

A thermogenic organ, modified from an eye muscle, warms the brain and eyes of several oceanic fish. The extraocular muscles associated with thermogenesis are composed of modified muscle cells that are structurally distinct from all other types of muscle previously described. In "heater" cells, contractile filaments are virtually absent and the cell volume is packed with mitochondria and smooth membranes. Freeze-fracture studies and negative staining of microsomal fractions treated with vanadate indicate that most of the membrane system of heater cells has a high Ca2+-ATPase density and is equivalent to skeletal muscle sarcoplasmic reticulum (SR). High voltage electron micrographs of heater cells infiltrated with the Golgi stain demonstrate that the cells also have an extensive transverse tubule system with a complicated three-dimensional structure. Junctional regions between transverse tubules and SR occur in the heater cell and contain feet protein. Activation of thermogenesis in heater cells may occur through the same protein components involved in excitation-contraction coupling and appears to be associated with the ATP-dependent cycling of calcium at the SR.

Characterization of the sarcoplasmic reticulum proteins in the thermogenic muscles of fish.

Marlins, sailfish, spearfishes, and swordfish have extraocular muscles that are modified into thermogenic organs beneath the brain. The modified muscle cells, called heater cells, lack organized myofibrils and are densely packed with sarcoplasmic reticulum (SR), transverse (T) tubules, and mitochondria. Thermogenesis in the modified extraocular muscle fibers is hypothesized to be associated with increased energy turnover due to Ca2+ cycling at the SR. In this study, the proteins associated with sequestering and releasing Ca2+ from the SR (ryanodine receptor, Ca2+ ATPase, calsequestrin) of striated muscle cells were characterized in the heater SR using immunoblot and immunofluorescent techniques. Immunoblot analysis with a monoclonal antibody that recognizes both isoforms of nonmammalian RYRs indicates that the fish heater cells express only the alpha RYR isoform. The calcium dependency of [3H]ryanodine binding to the RYR isoform expressed in heater indicates functional identity with the non-mammalian alpha RYR isoform. Fluorescent labeling demonstrates that the RYR is localized in an anastomosing network throughout the heater cell cytoplasm. Measurements of oxalate supported 45Ca2+ uptake, Ca2+ ATPase activity, and [32P]phosphoenzyme formation demonstrate that the SR contains a high capacity for Ca2+ uptake via an ATP dependent enzyme. Immunoblot analysis of calsequestrin revealed a significant amount of the Ca2+ binding protein in the heater cell SR. The present study provides the first direct evidence that the heater SR system contains the proteins necessary for Ca2+ release, re-uptake and sequestration, thus supporting the hypothesis that thermogenesis in the modified muscle cells is achieved via an ATP-dependent cycling of Ca2+ between the SR and cytosolic compartments.

Structural evidence for direct interaction between the molecular components of the transverse tubule/sarcoplasmic reticulum junction in skeletal muscle.

The architecture of the junctional sarcoplasmic reticulum (SR) and transverse tubule (T tubule) membranes and the morphology of the two major proteins isolated from these membranes, the ryanodine receptor (or foot protein) and the dihydropyridine receptor, have been examined in detail. Evidence for a direct interaction between the foot protein and a protein component of the junctional T tubule membrane is presented. Comparisons between freeze-fracture images of the junctional SR and rotary-shadowed images of isolated triads and of the isolated foot protein, show that the foot protein has two domains. One is the large hydrophilic foot which spans the junctional gap and is composed of four subunits. The other is a hydrophobic domain which presumably forms the SR Ca2+-release channel and which also has a fourfold symmetry. Freeze-fracture images of the junctional T tubule membranes demonstrate the presence of diamond-shaped clusters of particles that correspond exactly in position to the subunits of the feet protein. These results suggest the presence of a large junctional complex spanning the two junctional membranes and intervening gap. This junctional complex is an ideal candidate for a mechanical coupling hypothesis of excitation-contraction coupling at the triadic junction.

Evolution of endothermy in fish: mapping physiological traits on a molecular phylogeny.

Mackerels, tunas, and billfishes (suborder Scombroidei and Teleostei) provide an ideal taxonomic context in which to examine the evolution of endothermy. Multiple origins and diverse strategies for endothermy exist among these fish. Here a molecular phylogeny of the Scombroidei has been determined by direct sequencing of a portion of the mitochondrial cytochrome b gene. The distribution of endothermic species within this proposed genealogy indicates that the ability to warm the brain and retina arose independently in three lineages, each time in association with a movement into colder water. This suggests that the evolution of cranial endothermy in fish was selected in order to permit thermal niche expansion and not selected for increased aerobic capacity.

Migratory movements, depth preferences, and thermal biology of Atlantic bluefin tuna.

The deployment of electronic data storage tags that are surgically implanted or satellite-linked provides marine researchers with new ways to examine the movements, environmental preferences, and physiology of pelagic vertebrates. We report the results obtained from tagging of Atlantic bluefin tuna with implantable archival and pop-up satellite archival tags. The electronic tagging data provide insights into the seasonal movements and environmental preferences of this species. Bluefin tuna dive to depths of >1000 meters and maintain a warm body temperature. Western-tagged bluefin tuna make trans-Atlantic migrations and they frequent spawning grounds in the Gulf of Mexico and eastern Mediterranean. These data are critical for the future management and conservation of bluefin tuna in the Atlantic.

Habitat and behaviour of yellowfin tuna Thunnus albacares in the Gulf of Mexico determined using pop-up satellite archival tags.

This study presents the first data on movement, habitat use and behaviour for yellowfin tuna Thunnus albacares in the Atlantic Basin. Six individuals were tracked in the Gulf of Mexico using pop-up satellite archival tags. Records up to 80 days in length were obtained, providing information on depth and temperature preferences as well as horizontal movements. Thunnus albacares in the Gulf of Mexico showed a strong preference for the mixed layer and thermocline, consistent with findings for this species in other ocean basins. Fish showed a diel pattern in depth distribution, remaining in surface and mixed layer waters at night and diving to deeper waters during the day. The vertical extent of T. albacares habitat appeared to be temperature limited, with fish generally avoiding waters that were >6 degrees C cooler than surface waters. The vertical and thermal habitat usage of T. albacares differs from that of bigeye Thunnus obesus and bluefin Thunnus thynnus, Thunnus orientalis and Thunnus maccoyii tunas. These results are consistent with the results of earlier studies conducted on T. albacares in other oceans.

Surface plasmon induced polarization rotation and optical vorticity in a single mode waveguide.

The control and manipulation of the mode polarization state in a single mode dielectric waveguide is of considerable significance for optical information processing utilizing the polarization state to store digital information and integrated photonic devices used for high speed signaling. Here we report on an integrated on-chip mode polarization rotation based on short metal Cu electrodes placed in close proximity to the dielectric waveguide core. Polarization mode rotation with specific rotation of 10(4) degrees/mm is observed for offset metallic electrodes placed diagonally along a single mode dielectric waveguide. The mechanism for the polarization rotation is shown to be directional coupling into guided surface plasmon modes at the metal corners and coupling between the guided plasmon modes. This inter-plasmon coupling gives rise to giant polarization rotation and optical vorticity (helical power flow) in the waveguide.

Ion permeation and block of M-type and delayed rectifier potassium channels. Whole-cell recordings from bullfrog sympathetic neurons.

Ion permeation and conduction were studied using whole-cell recordings of the M-current (I(M)) and delayed rectifier (IDR), two K+ currents that differ greatly in kinetics and modulation. Currents were recorded from isolated bullfrog sympathetic neurons with 88 mM [K+]i and various external cations. Selectivity for extracellular monovalent cations was assessed from permeability ratios calculated from reversal potentials and from chord conductances for inward current. PRb/PK was near 1.0 for both channels, and GRb/GK was 0.87 +/- 0.01 for IDR but only 0.35 +/- 0.01 for I(M) (15 mM [Rb+]o or [K+]o). The permeability sequences were generally similar for I(M) and IDR: K+ approximately Rb+ > NH4+ > Cs+, with no measurable permeability to Li+ or CH3NH3+. However, Na+ carried detectable inward current for IDR but not I(M). Nao+ also blocked inward K+ current for IDR (but not IM), at an apparent electrical distance (delta) approximately 0.4, with extrapolated dissociation constant (KD) approximately 1 M at 0 mV. Much of the instantaneous rectification of IDR in physiologic ionic conditions resulted from block by Nao+. Extracellular Cs+ carried detectable inward current for both channel types, and blocked I(M) with higher affinity (KD = 97 mM at 0 mV for I(M), KD) approximately 0.2 M at 0 mV for IDR), with delta approximately 0.9 for both. IDR showed several characteristics reflecting a multi-ion pore, including a small anomalous mole fraction effect for PRb/PK, concentration-dependent GRb/GK, and concentration-dependent apparent KD's and delta's for block by Nao+ and Cso+. I(M) showed no clear evidence of multi-ion pore behavior. For I(M), a two-barrier one-site model could describe permeation of K+ and Rb+ and block by Cso+, whereas for IDR even a three-barrier, two-site model was not fully adequate.