Ultra-High Foraging Rates of Harbor Porpoises Make Them Vulnerable to Anthropogenic Disturbance.

The question of how individuals acquire and allocate resources to maximize fitness is central in evolutionary ecology. Basic information on prey selection, search effort, and capture rates are critical for understanding a predator's role in its ecosystem and for predicting its response to natural and anthropogenic disturbance. Yet, for most marine species, foraging interactions cannot be observed directly. The high costs of thermoregulation in water require that small marine mammals have elevated energy intakes compared to similar-sized terrestrial mammals [1]. The combination of high food requirements and their position at the apex of most marine food webs may make small marine mammals particularly vulnerable to changes within the ecosystem [2-4], but the lack of detailed information about their foraging behavior often precludes an informed conservation effort. Here, we use high-resolution movement and prey echo recording tags on five wild harbor porpoises to examine foraging interactions in one of the most metabolically challenged cetacean species. We report that porpoises forage nearly continuously day and night, attempting to capture up to 550 small (3-10 cm) fish prey per hour with a remarkable prey capture success rate of >90%. Porpoises therefore target fish that are smaller than those of commercial interest, but must forage almost continually to meet their metabolic demands with such small prey, leaving little margin for compensation. Thus, for these "aquatic shrews," even a moderate level of anthropogenic disturbance in the busy shallow waters they share with humans may have severe fitness consequences at individual and population levels.

Quantitative Measures of Anthropogenic Noise on Harbor Porpoises: Testing the Reliability of Acoustic Tag Recordings.

In recent years, several sound and movement recording tags have been developed to sample the acoustic field experienced by cetaceans and their reactions to it. However, little is known about how tag placement and an animal's orientation in the sound field affect the reliability of on-animal recordings as proxies for actual exposure. Here, we quantify sound exposure levels recorded with a DTAG-3 tag on a captive harbor porpoise exposed to vessel noise in a controlled acoustic environment. Results show that flow noise is limiting onboard noise recordings, whereas no evidence of body shading has been found for frequencies of 2-20 kHz.

Echolocation by the harbour porpoise: life in coastal waters.

The harbor porpoise is one of the smallest and most widely spread of all toothed whales. They are found abundantly in coastal waters all around the northern hemisphere. They are among the 11 species known to use high frequency sonar of relative narrow bandwidth. Their narrow biosonar beam helps isolate echoes from prey among those from unwanted items and noise. Obtaining echoes from small objects like net mesh, net floats, and small prey is facilitated by the very high peak frequency around 130 kHz with a wavelength of about 12 mm. We argue that such echolocation signals and narrow band auditory filters give the harbor porpoise a selective advantage in a coastal environment. Predation by killer whales and a minimum noise region in the ocean around 130 kHz may have provided selection pressures for using narrow bandwidth high frequency biosonar signals.

High source levels and small active space of high-pitched song in bowhead whales (Balaena mysticetus).

The low-frequency, powerful vocalizations of blue and fin whales may potentially be detected by conspecifics across entire ocean basins. In contrast, humpback and bowhead whales produce equally powerful, but more complex broadband vocalizations composed of higher frequencies that suffer from higher attenuation. Here we evaluate the active space of high frequency song notes of bowhead whales (Balaena mysticetus) in Western Greenland using measurements of song source levels and ambient noise. Four independent, GPS-synchronized hydrophones were deployed through holes in the ice to localize vocalizing bowhead whales, estimate source levels and measure ambient noise. The song had a mean apparent source level of 185±2 dB rms re 1 µPa @ 1 m and a high mean centroid frequency of 444±48 Hz. Using measured ambient noise levels in the area and Arctic sound spreading models, the estimated active space of these song notes is between 40 and 130 km, an order of magnitude smaller than the estimated active space of low frequency blue and fin whale songs produced at similar source levels and for similar noise conditions. We propose that bowhead whales spatially compensate for their smaller communication range through mating aggregations that co-evolved with broadband song to form a complex and dynamic acoustically mediated sexual display.

Seasonal changes in the vocal behavior of bowhead whales (Balaena mysticetus) in Disko Bay, Western-Greenland.

Singing behavior has been described from bowhead whales in the Bering Sea during their annual spring migration and from Davis Strait during their spring feeding season. It has been suggested that this spring singing behavior is a remnant of the singing during the winter breeding season, though no winter recordings are available. In this study, the authors describe recordings made during the winter and spring months of bowhead whales in Disko Bay, Western-Greenland. A total of 7091 bowhead whale sounds were analyzed to describe the vocal repertoire, the singing behavior, and the changes in vocal behavior from February to May. The vocal signals could be divided into simple (frequency-modulated) calls (n=483), complex (amplitude-modulated) calls (n=635), and song notes (n=5973). Recordings from the end of February to middle of March were characterized by higher call rates with a greater diversity of call types than recordings made later in the season. This study is the first description of bowhead song from the stock in Western-Greenland during both the winter and spring months, and provides support for the hypothesis that song during the winter months contains more song notes than song from the spring making the winter song more variable.

Underwater noise from three types of offshore wind turbines: estimation of impact zones for harbor porpoises and harbor seals.

Underwater noise was recorded from three different types of wind turbines in Denmark and Sweden (Middelgrunden, Vindeby, and Bockstigen-Valar) during normal operation. Wind turbine noise was only measurable above ambient noise at frequencies below 500 Hz. Total sound pressure level was in the range 109-127 dB re 1 microPa rms, measured at distances between 14 and 20 m from the foundations. The 1/3-octave noise levels were compared with audiograms of harbor seals and harbor porpoises. Maximum 1/3-octave levels were in the range 106-126 dB re 1 microPa rms. Maximum range of audibility was estimated under two extreme assumptions on transmission loss (3 and 9 dB per doubling of distance, respectively). Audibility was low for harbor porpoises extending 20-70 m from the foundation, whereas audibility for harbor seals ranged from less than 100 m to several kilometers. Behavioral reactions of porpoises to the noise appear unlikely except if they are very close to the foundations. However, behavioral reactions from seals cannot be excluded up to distances of a few hundred meters. It is unlikely that the noise reaches dangerous levels at any distance from the turbines and the noise is considered incapable of masking acoustic communication by seals and porpoises.

Echolocation by two foraging harbour porpoises (Phocoena phocoena).

Synchronized video and high-frequency audio recordings of two trained harbour porpoises searching for and capturing live fish were used to study swimming and echolocation behaviour. One animal repeated the tasks blindfolded. A splash generated by the fish being thrown into the pool or - in controls - by a boat hook indicated prey and stimulated search behaviour. The echolocation sequences were divided into search and approach phases. In the search phase the porpoises displayed a clear range-locking behaviour on landmarks, indicated by a distance-dependent decrease in click interval. Only in trials with fish was the search phase followed by an approach phase. In the initial part of the approach phase the porpoises used a rather constant click interval of around 50 ms. The terminal part started with a sudden drop in click interval at distances around 2-4 m. Close to the prey the terminal part ended with a buzz, characterized by constant click intervals around 1.5 ms. The lag time in the search and the initial part of the approach phase seems to be long enough for the porpoise to process echo information before emitting the next click (pulse mode). However, we assume that during the buzz lag times are too short for pulse mode processing and that distance information is perceived as a ;pitch' with a ;frequency' corresponding to the inverse of the two-way transit time (pitch mode). The swimming speed of the animal was halved when it was blindfolded, while the click intervals hardly changed, resulting in more clicks emitted per metre swum.

Auditory temporal resolution of a wild white-beaked dolphin (Lagenorhynchus albirostris).

Adequate temporal resolution is required across taxa to properly utilize amplitude modulated acoustic signals. Among mammals, odontocete marine mammals are considered to have relatively high temporal resolution, which is a selective advantage when processing fast traveling underwater sound. However, multiple methods used to estimate auditory temporal resolution have left comparisons among odontocetes and other mammals somewhat vague. Here we present the estimated auditory temporal resolution of an adult male white-beaked dolphin, (Lagenorhynchus albirostris), using auditory evoked potentials and click stimuli. Ours is the first of such studies performed on a wild dolphin in a capture-and-release scenario. The white-beaked dolphin followed rhythmic clicks up to a rate of approximately 1,125-1,250 Hz, after which the modulation rate transfer function (MRTF) cut-off steeply. However, 10% of the maximum response was still found at 1,450 Hz indicating high temporal resolution. The MRTF was similar in shape and bandwidth to that of other odontocetes. The estimated maximal temporal resolution of white-beaked dolphins and other odontocetes was approximately twice that of pinnipeds and manatees, and more than ten-times faster than humans and gerbils. The exceptionally high temporal resolution abilities of odontocetes are likely due primarily to echolocation capabilities that require rapid processing of acoustic cues.

The utility of cyclodextrins for enhancing oral bioavailability.

Cyclodextrins (CD) have been utilized extensively in pharmaceutical formulations to enhance oral bioavailability. A critical review of the literature in which cyclodextrins were utilized for this purpose was conducted. The goal of this review was to determine if quantitative guidelines for drug and cyclodextrin properties necessary for bioavailability enhancement using cyclodextrins could be extracted. Twenty-eight studies were examined in which the focus was on the use of cyclodextrins as solubilizers to enhance bioavailability. Commonly observed factors included: utilization of pre-formed complex rather than physical mixtures, drug hydrophobicity (logP > 2.5), low drug solubility (typically< 1 mg/ml), moderate binding constant (< 5000 M(-1)), low dose (< 100 mg), and low CD:drug ratio (< 2:1). These general guidelines, however, did not apply to all studies. Quantitative guidelines useful to a formulation scientist considering the use of cyclodextrins were difficult to develop due to missing information and the complicated manner in which drug and cyclodextrin properties interact to influence key drug delivery processes (e.g., dissolution, absorption). The mechanisms by which cyclodextrins influence these processes, again emphasizing solubilization capabilities, are discussed to provide further insight into why cyclodextrins will increase bioavailability in certain cases but not influence or possibly decrease bioavailability in others.

Echolocation clicks from killer whales (Orcinus orca) feeding on herring (Clupea harengus).

Echolocation clicks from Norwegian killer whales feeding on herring schools were recorded using a four-hydrophone array. The clicks had broadband bimodal frequency spectra with low and high frequency peaks at 24 and 108 kHz, respectively. The -10 dB bandwidth was 35 kHz. The average source level varied from 173 to 202 dB re 1 microPa (peak-to-peak) at 1 m. This is considerably lower than source levels described for Canadian killer whales foraging on salmon. It is suggested that biosonar clicks of Norwegian killer whales are adapted for localization of prey with high target strength and acute hearing abilities.

Practical considerations in development of solid dosage forms that contain cyclodextrin.

The following is a review of the literature that addresses the use of cyclodextrin in solid dosage forms. Care was taken to exclude physical and chemical characteristics of cyclodextrin, which have been discussed in the literature. A flow diagram is provided to outline the decision-making steps that are involved in the development process. Both preparation of physical mixtures and inclusion complexes are considered. Analytical techniques to determine the presence of inclusion complexes, the effect of other excipients on complex formation, the effect of size limitation of solid dosages forms, powder processing, and storage of solid dosage forms are discussed.

Mechanistic investigation of drug release from asymmetric membrane tablets: effect of media gradients (osmotic pressure and concentration), and potential coating failures on in vitro release.

PURPOSE: An asymmetric membrane (AM) tablet was developed for a soluble model compound to study the in vitro drug release mechanisms in challenge conditions, including osmotic gradients, concentration gradients, and under potential coating failure modes. Porous, semipermable membrane integrity may be compromised by a high fat meal or by the presence of a defect in the coating that could cause a safety concern about dose-dumping. METHODS: The osmotic and diffusional release mechanisms of the AM tablet were independently shut down such that their individual contribution to the overall drug release was measured. Shut off of osmotic and diffusional release was accomplished by performing dissolution studies into receptor solutions with osmotic pressure above the internal core osmotic pressure and into receptor solutions saturated with drug, respectively. The effect of coating failure modes on in vitro drug release from the AM tablet was assessed through a simulated high-fat meal and by intentionally compromising the coating integrity. RESULTS: The predominant drug release mechanism for the AM tablet was osmotic and accounted for approximately 90-95% of the total release. Osmotic release was shutoff when the receptor media osmotic pressure exceeded 76 atm. Diffusional release of the soluble drug amounted to 5-10% of the total release mechanism. The observed negative in vitro food effect was attributed to the increased osmotic pressure from the high fat meal when compared to the predicted release rates in sucrose media with the same osmotic pressure. This suppression in drug release rate due to a high fat meal is not anticipated to affect in vivo performance of the dosage form, as the rise in pressure is short-lived. CONCLUSIONS: Drug release from the AM system studied was determined to be robust to varying and extreme challenge conditions. The conditions investigated included varying pH, agitation rate, media osmotic pressure, media saturated with drug to eliminate the concentration gradient, simulated high fat meal, and intentionally placed film coating defects. Osmotic and diffusional shut off experiments suggest that the mechanism governing drug release is a combination of osmotic and diffusional at approximately 90-95% and 5-10%, respectively. In addition, the coating failure mode studies revealed this formulation and design is not significantly affected by a high fat meal or by an intentionally placed defect in the film coating, and more specifically, did not result in a burst of drug release.

Auditory brainstem response in a harbor porpoise show lack of automatic gain control for simulated echoes.

The auditory brainstem response (ABR) response to simulated echolocation clicks was studied in a harbor porpoise, Phocoena phocoena, to determine the relationship between the animal's perceived echo strength and the simulated target distance. In one experiment the click level at the listening post was kept constant while delay was changed, in another, the level was varied to approximate spreading losses. Results of both experiments indicate that there is no automatic gain control in the hearing system of this harbor porpoise.

Spatial orientation in echolocating harbour porpoises (Phocoena phocoena).

Studies concerning the echolocation behaviour of odontocetes focus mainly on target detection and discrimination, either in stationary animals or in animals approaching a specific target. We present the first data on the use of echolocation for spatial orientation or navigation. Synchronised video and high-frequency recordings were made of two harbour porpoises trained to swim from one position to another across an outdoor pool in order to correlate swimming and echolocation behaviour. Both porpoises showed a clear range-locking behaviour on specific positions near the end of the pool, as indicated by a decrease in click interval with decreasing distance. The decrease in click interval followed the two-way-transit time, which is the time interval between the outgoing click and the received echo from the focal object. This suggests that the porpoises used focal objects as landmarks. The lag time, defined as the time between the arrival of an echo from a landmark and the emission of the next click, was task specific. The lag time was longer for difficult tasks (26-36 ms) and shorter for simpler tasks (14-19 ms), with some individual differences between the two animals. Our results suggest that echolocation by odontocetes is used not only for target detection, localisation and classification but also for spatial orientation.

Acoustic characteristics of underwater tail slaps used by Norwegian and Icelandic killer whales (Orcinus orca) to debilitate herring (Clupea harengus).

Norwegian killer whales debilitate prey by slapping their tails into herring schools. These underwater tail slaps produce a thud-like sound. It is unclear whether this sound is caused by cavitation and/or physical contact between herring and whale tail. Also the forces causing debilitation of the fish are not understood. Here we present an acoustic analysis of underwater tail slaps using a multi-channel wide (150 kHz) band recording system. Underwater tail slaps produced by Norwegian killer whales generated sounds consisting of multiple pulses with source levels of 186+/-5.4 dB (pp) re.1 microPa at 1 m (+/-1 s.d., N = 4). The -3 dB and 97% energy bandwidths were 36.8+/-22.5 kHz and 130.5+/-17.5 kHz (+/-1 s.d., N = 13), respectively, with a centre frequency of 46.1+/-22.3 kHz. The similarities between the acoustic properties of underwater tail slaps recorded from killer whales in Norway, and thud-like sounds recorded from killer whales in Iceland suggest that Norwegian and Icelandic killer whales use similar hunting techniques. The acoustic characteristics of sounds produced by underwater tail slaps were similar to the ones from other cavitation sound sources described in the literature, both in term of temporal and frequency features as well as in source level. We suggest that multiple factors generated by the tail slaps like particle fluctuations, turbulence, pressure changes and physical impact cause debilitation of herring.

Estimated transmission beam pattern of clicks recorded from free-ranging white-beaked dolphins (Lagenorhynchus albirostris).

Recordings were made from white-beaked dolphins in Icelandic waters using a four-hydrophone array in a star configuration. The acoustic signals were amplified and sampled to a hard disk at a rate of 800 kHz per channel. The 3 and 10 dB beamwidths were calculated to be 8 degrees and 10 degrees, respectively, indicating a narrower transmission beam for white-beaked dolphins than that reported for bottlenose dolphins (Tursiops truncatus). The beamwidth was more similar to that found for belugas (Delphinapterus lucas). The measured beam pattern included large side lobes, perhaps due to the inclusion of off-axis clicks, even after applying several criteria to select only on-axis clicks. The directivity index was calculated to be 18 dB when using all data for angles from 0 degrees-50 degrees. The calculated sound radiation from a circular piston with a radius of 6 cm driven by a white-beaked dolphin click had a beam pattern very similar to the measured beam pattern for the main transmission lobe of the white-beaked dolphin. The directivity index was 29 dB. This is the first attempt to estimate the directionality index of dolphins in the field.