Whales Unit Study

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Odontocetes produce rapid bursts of high-frequency clicks that are thought to be primarily for echolocation. Specialized organs in an odontocete produce collections of clicks and buzzes at frequencies from 0. Lower frequencies are used for distance echolocation, due to the fact that shorter wavelengths do not travel as far as longer wavelengths underwater.

Higher frequencies are more effective at shorter distances, and can reveal more detailed information about a target. Echoes from clicks convey, not only the distance to the target, but also the size, shape, speed, and vector of its movement. Additionally, echolocation allows the odontocete to easily discern the difference between objects that are different in material composition, even if visually identical, by their different densities.

Individuals also appear to be able to isolate their own echoes during pod feeding activity without interference from other pod members' echolocations. Whistles are used for communication, and four- to six-month-old calves develop unique sounds that they use most frequently throughout their lives. Such "signature whistles" are distinctive to the individual and may serve as a form of identification among other odontocetes. Frankel quotes one researcher who says listening to a school of odontocetes is like listening to a group of children at a school playground.

The multiple sounds odontocetes make are produced by passing air through a structure in the head called the phonic lips. As the air passes through this narrow passage, the phonic lip membranes are sucked together, causing the surrounding tissue to vibrate. These vibrations can, as with the vibrations in the human larynx, be consciously controlled with great sensitivity.

Every toothed whale except the sperm whale has two sets of phonic lips and is thus capable of making two sounds independently. From there, the air may be recycled back into the lower part of the nasal complex, ready to be used for sound creation again, or passed out through the blowhole. The French name for phonic lips, museau de singe , translates literally as "monkey's muzzle", which the phonic lip structure is supposed to resemble. Baleen whales formally called mysticetes do not have phonic lip structure.

Instead, they have a larynx that appears to play a role in sound production, but it lacks vocal cords, and scientists remain uncertain as to the exact mechanism. It is likely that they recycle air around the body for this purpose. There are at least nine separate blue whale acoustic populations worldwide. Calls are progressively getting lower in frequency. For example the Australian pygmy blue whales are decreasing their mean call frequency rate at approximately 0.

The migration patterns of blue whales remains unclear.

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Some populations appear to be resident in habitats of year-round high productivity in some years, [17] while others undertake long migrations to high-latitude feeding grounds, but the extent of migrations and the components of the populations that undertake them are poorly known. While the complex sounds of the humpback whale and some blue whales are believed to be primarily used in sexual selection , [21] the simpler sounds of other whales have a year-round use. For instance, the depth of water or the existence of a large obstruction ahead may be detected by loud noises made by baleen whales.

The question of whether whales sometimes sing purely for aesthetic enjoyment, personal satisfaction, or 'for art's sake', is considered by some to be "an untestable question". Two groups of whales, the humpback whale and the subspecies of blue whale found in the Indian Ocean , are known to produce a series of repetitious sounds at varying frequencies known as whale song.

Marine biologist Philip Clapham describes the song as "probably the most complex in the animal kingdom. Male humpback whales perform these vocalizations often during the mating season, and so it is believed the purpose of songs is to aid mate selection. Singing has also been recorded in competitive groups of whales that are composed of one female and multiple males. Interest in whale song was aroused by researchers Roger Payne and Scott McVay after the songs were brought to their attention by a Bermudian named Frank Watlington who was working for the US government at the SOFAR station listening for Russian submarines with underwater hydrophones off the coast of the island.

Preschool Whale Unit

The songs follow a distinct hierarchical structure. The base units of the song sometimes loosely called the " notes " are single uninterrupted emissions of sound that last up to a few seconds. The units may be frequency modulated i. However, the adjustment of bandwidth on a spectrogram representation of the song reveals the essentially pulsed nature of the FM sounds. A collection of four or six units is known as a sub- phrase , lasting perhaps ten seconds see also phrase music. A whale will typically repeat the same phrase over and over for two to four minutes.

This is known as a theme. A collection of themes is known as a song. All the whales in an area sing virtually the same song at any point in time and the song is constantly and slowly evolving over time. The pace of evolution of a whale's song also changes—some years the song may change quite rapidly, whereas in other years little variation may be recorded. Whales occupying the same geographical areas which can be as large as entire ocean basins tend to sing similar songs, with only slight variations.

Whales from non-overlapping regions sing entirely different songs. As the song evolves, it appears that old patterns are not revisited. Humpback whales may also make stand-alone sounds that do not form part of a song, particularly during courtship rituals. Humpbacks generally feed cooperatively by gathering in groups, swimming underneath shoals of fish and all lunging up vertically through the fish and out of the water together.

Prior to these lunges, whales make their feeding call. The exact purpose of the call is not known, but research suggests that fish know what it means. Some scientists have proposed that humpback whale songs may serve an echolocative purpose, [29] but this has been subject to disagreement.

Humpback whales have also been found to make a range of other social sounds to communicate such as "grunts", "groans", "thwops", "snorts" and "barks". Most baleen whales make sounds at about 15—20 hertz. There is disagreement in the scientific community regarding the uniqueness of the whale's vocalization [34] and whether it is a member of a hybrid whale [34] such as the well documented Blue and Fin Whale hybrids. In , researchers found that blue whale song has been deepening in its tonal frequency since the s.

Short range calls are reported during social and resting periods while long range are more commonly reported during foraging and feeding. Most other whales and dolphins produce sounds of varying degrees of complexity. Of particular interest is the Beluga the "sea canary" which produces an immense variety of whistles, clicks and pulses.

Researchers use hydrophones often adapted from their original military use in tracking submarines to ascertain the exact location of the origin of whale noises. Christopher Clark of Cornell University conducted using military data showed that whale noises travel for thousands of kilometres. An important finding is that whales, in a process called the Lombard effect , adjust their song to compensate for background noise pollution.

Prior to the introduction of human noise production, Clark says the noises may have travelled right from one side of an ocean to the other, agreeing with a thirty-year-old concept blaming large-scale shipping. Environmentalists fear that such boat activity is putting undue stress on the animals as well as making it difficult to find a mate. In the past decade, many effective automated methods, such as signal processing, data mining, and machine learning techniques have been developed to detect and classify whale vocalizations.

Kelly was the first person known to recognize whale singing for what it was, while on the brig Eliza in the Sea of Japan in From Wikipedia, the free encyclopedia. Recording of humpback whales singing and clicking. List of whale vocalizations. A humpback whale song. However, blow analysis is still in its infancy, and many validation questions remain to be addressed.

Sampling methodology will require detailed study, e. Continued collaborations between human breath researchers and cetacean conservation biologists will be most useful.

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Since the early s, dart biopsying has become one of the most common collection methods for obtaining biological tissue samples from free-ranging cetaceans Lambertsen, ; Mathews et al. These samples have traditionally provided information about diet lipid composition and stable isotopes , pollutant exposure, and genetics Noren and Mocklin, Increasingly, they are also being used to study the physiological states of sampled individuals, by measuring lipophilic hormones especially steroids in the blubber and by characterizing gene expression in the epidermis.

Dart biopsy samples are predominantly collected using either a crossbow or a pneumatic rifle with modified dart tips Lambertsen, ; Mathews et al. Here, we will use the terms epidermis and blubber to distinguish these two layers. It may also be possible to collect epidermal samples in the form of sloughed skin, which is occasionally visible at the water surface after vigorous activites, such as breeching. Such samples have been collected successfully for genetic analyses e. Recently, a nascent but growing field of research has focused on measurement of lipophilic hormones, especially steroids, in blubber Mansour et al.

Typically, the tissue has been processed using a multistep organic solvent extraction to isolate lipophilic hormones, and then these hormones are measured using immunoassays or chromatography separation followed by mass spectrometry. Several investigators have used immunoassay of blubber progesterone to assess pregnancy state in several delphinids, Kellar et al. These studies have shown large differences in blubber progesterone of pregnant vs. Likewise, in short-beaked common dolphins Delphinus delphis during the presumed mating period, sexually mature males had substantially higher levels of blubber testosterone than immature males, with no overlap between the two groups Kellar et al.

Outside the mating period, mature males still had significantly higher testosterone than immature animals, but with more overlap between groups. Thus, blubber hormone analyses may prove to be a useful complement to faecal hormone and blow hormone analyses. Other lipophilic hormones are also likely to be measurable in blubber, including cortisol, aldosterone, and thyroid hormones Kershaw and Flier, ; MacKenzie et al.

Blubber hormone analysis is still relatively novel.

Next steps will involve applying the techniques to more species particularly mysticetes , and increasing the number of lipophilic hormones validated. As with faecal and blow hormone analyses, validations for large whales will need to include a phase of physiological validations, i. In addition, little is known about the temporal dynamics of hormone deposition in cetacean blubber, i.

Likewise, there may also be lag between reduced levels in blood and clearance from blubber. Blubber biopsies have long been used to measure exposure to organic pollutants. Given that cetaceans are long-lived predators, they are predisposed to the accumulation of lipophilic and bioaccumulative contaminants, such as persistent organic pollutants POPs. Persistent organic pollutants are a significant concern for cetacean health and population sustainability Tanabe et al.

Most biomonitoring efforts rely on blubber biopsies to assess contaminant exposure in wild cetacean populations, because the blubber is the primary site of accumulation for lipophilic contaminants Tanabe et al. Extensive studies have been carried out using blubber biopsy samples to assess the contaminant burdens of a number of cetacean species and populations Tanabe et al. Overall, males tend to have higher POP concentrations in blubber than females, because females rid themselves of some of their POP burden during lactation.

Pollutant levels are positively correlated with increasing trophic level and negatively correlated with body size Borrell, Age and sex influences on accumulation must be taken into consideration Tuerk et al. Generally, analytical procedures for contaminant studies tend to be expensive and time consuming, but the information produced has been invaluable for assessing contaminant exposure and consequent effects on reproduction, mortality, immune function, and other parameters relevant to conservation physiology.

Blubber contains a variety of different fatty acids FAs , and the types and proportions of FAs present can reveal information relevant to dietary physiology, thermal physiology, and even the age of the individual. Fatty acid profiles have been used to answer qualitative questions about spatial or temporal variation in diets between individuals or populations Budge et al. Fatty acid analysis has been used to assess stock structure, trophic positions, diet, and individual age in a variety of mysticetes and odontocetes Borobia et al.

One complication is that cetacean blubber is highly stratified Koopman et al. Finally, variations in FA characteristics are known to affect insulative properties of the blubber and thus can be used to study aspects of thermal physiology pygmy sperm whales, Kogia breviceps , and short-finned pilot whales, Globicephala macrorhynchus , Bagge et al.

Epidermal diseases in free-ranging whales and dolphins have been studied primarily using photographic analyses see next main section , but can also be studied via epidermal and blubber samples. Current understanding of cetacean epidermal diseases is based on samples taken from stranded and bycaught animals Baker, ; Van Bressem et al. Where possible, collection of biopsy samples from lesions of free-ranging animals would increase our knowledge of pathogens affecting cetaceans and contribute to a better understanding of the pathogenesis of epidermal lesions.

A unique aspect of biopsy samples, in comparison to other matrices faeces and blow , is that biopsy samples contain large quantities of intact peptides and proteins. In terrestrial mammals, proteome-associated profiling is a useful tool to assess energetic balance, immune system function, contaminant exposure, and responses to a diverse range of environmental stressors Silvestre et al. A large variety of identification and quantification methods are available for proteomics studies; these include liquid chromatography and two-dimensional polyacrylamide gel electrophoresis Brewis and Brennan, , tandem mass spectrometry Dhingra et al.

There is, as yet, limited molecular information for many large whales, which limits applications of certain species-specific proteomics tools Veldhoen et al. However, those proteins and peptides that appear to be highly conserved across mammals could probably be investigated in large whales with relative ease. Preliminary studies in small odontocetes over the past decade have been encouraging, and indicate that proteomics and transcriptomics may be useful for the types of physiological information outlined below. Post-mortem liver analyses of sperm whales Boon et al. It is now known that these and related enzymes also occur in the endothelium of the dermal papillae Bickers et al.

These enzymes have now been measured in epidermal samples from over 17 species of cetaceans, including both odontocetes and mysticetes Fossi et al. These changes, in turn, induce counteracting molecular stress responses, including expression of SRPs that are detectable in many tissues and organs, including the epidermis Arck et al. The analytical approach involves assessment of molecular profiles in the epidermis that are indicative of a stress response. For example, spotted dolphins Stenella attenuata subjected to acute stress associated with chase and purse-seine encirclement exhibited altered SRP expression in epidermal samples, in comparison to control dolphins Dizon et al.

The spotted dolphin studies indicate that this method may be suitable for assessing chronic stress e. Additionally, the SRP analytical technique is rapid and relatively inexpensive. The SRPs used in these dolphin studies are thought to be highly conserved across mammals and are likely to be present in large whales as well. Many validations still remain to be done, including age, sex, and stressor relationships in different species, duration of the signal, possible decreased responsiveness to frequently repeated stressors Southern et al.

Transcriptomics has become a widely used approach for the study of normal and diseased gene expression in human skin Cole et al. Some transcriptome studies on cetacean epidermis have been conducted in vitro Ellis et al. Methods have also been published for sample collection and the development of cetacean cell lines Marsili et al. For example, Romano and Warr have developed genomics-based diagnostic methods for health assessment of bottlenose dolphins Tursiops truncatus using microarray techniques based on blood-based biomarkers related to immune response and stress.

However, application of these microarrays to biopsy samples awaits an assessment of the degree to which these genes are also expressed in dermis and epidermis. Various in vivo and in vitro studies of dolphins have also investigated altered gene expression profiles in response to contaminant exposure Mollenhauer et al. In summary, biopsy samples contain a wide variety of invaluable physiological information, ranging from lipophilic hormones and contaminants to CYP-related enzymes, stress-related proteins, RNA and associated indices of gene expression, and potential information on disease states.

Second, sampling rate is high—to date, higher than for faeces or blow collection—especially when biopsying effort is coupled with traditional line-transect surveying. In fact, several previous studies have resulted in over samples collected during a single survey season e. One disadvantage of dart biopsying is that it is invasive e. Fortunately, the small wounds generated by biopsy darting appear to be minor Noren and Mocklin, , and behavioural responses are minimal Gauthier and Sears, ; Clapham and Mattila, Genome sequences for a few marine mammal species are beginning to emerge, but coverage is still limited.

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One solution has been to use sequence databases from phylogenetically similar species, thereby allowing identification of highly conserved proteins. As discussed above, there has been parallel interest for developing similar techniques proteomics, transcriptomics, and genomics for blow and faecal samples. Such new and emerging molecular approaches will benefit from interdisciplinary research collaborations involving both cetacean field biologists and researchers from other disciplines.

Visual assessment of the external appearance of an animal is a time-honoured technique for assessing the health and nutritional status of individuals. Subjective assessment has recently been augmented by a variety of semi-quantitative and quantitative methods, which allow more rigorous assessment of changes in large whale body condition, within and between years, in the context of varying food supplies and other conservation-related stressors. The basic premise is that blubber, muscle, and visceral mass increase with nutritional status, enhancing the ability of females to conceive, suckle, and wean offspring, and also enhancing reproductive success of males, and that these changes in mass can be detected by assessment of overall body shape.

Furthermore, particular visible aspects, such as skin condition, parasite load and distribution, fresh wounds, and old scars from vessel and fishing gear interactions can contribute additional information on health. In small marine mammals, such health assessment can be undertaken by capture and release Wells et al. Most imaging techniques to assess appearance have been based on photographs from planes Fig.

Boat-based photography produces close-up lateral views of whatever portion of the animal is above the surface of the water typically, the dorsal surface of the animal, and often the entire tail when the animal dives. Aerial photographs are necessarily taken from a greater distance, but if obtained from a perpendicular vantage point above the animal, they provide a unique whole-body perspective that enables measurement of body length-to-width ratios, and true calibrated measurements if altitude and scale are available.

Finally, infrared thermography can potentially provide additional information relevant to thermal physiology. This image of an entangled, emaciated North Atlantic right whale Eubalaena glacialis ; Eg has been marked for length-to-width ratio analysis for comparison to unentangled animals, to assess likely body weight prior to dosing with sedatives for disentanglement efforts van der Hoop et al.

These visual health-assessment photographs show three North Atlantic right whales Eubalaena glacialis in good a , fair b and poor body condition c. Body condition is evaluated based on the degree of convexity good or concavity fair or poor in the dorsal back profile in the post-blowhole area white arrows. The whale in poor body condition c is entangled in fishing line and has other indicators of severely compromised health, including white skin lesions ellipses and rake marks forward of the blowholes rectangle.

Photos a and b: Initial aerial measurement of whales at sea primarily focused on measuring body length Best and Ruther, ; Angliss et al. Body length is often used as a proxy for age in cetacean biology, and many age-related aspects of whale physiology are traditionally expressed in terms of body length e. More recently, measurements of length in association with widths in NARWs have allowed estimates of body volume in addition to length Miller et al. Loss of body mass was also observed during lactation.

Thus, body width provides a useful measure of nutritional status Miller et al.

Whale vocalization

An integration of these body condition measures has been obtained by quantifying transitions between swimming and gliding for tagged diving whales. On the ascent, gliding begins once lung expansion overcomes the negative buoyancy of the rest of the animal. Thus, thinner whales, with less fat, will start to glide later during the ascent than fatter animals. Given that body fat can affect diving characteristics, with associated metabolic costs, measures of body condition are highly relevant to many energetic and physiological studies.

Lateral body condition assessment from a boat is only semi-quantitative, given the inability to observe the full submerged body outline, but data indicate that viewing even a portion of the animal may provide useful information on individual health. For example, boat-based photography allows a close-up view of skin condition with a level of detail that is not visible from manned aeroplanes Hamilton and Marx, This method uses a scoring system to evaluate body condition using the following parameters: Comparison of body condition scores of females during calving and non-calving years indicated that females had poorer body condition in calving years, and also in the year after calving compared with the year before calving.

Animals that later disappeared e. Comparison of these body condition scores to blubber thicknesses measured acoustically at sea Moore et al. An extension of this method used on western Pacific grey whales Bradford et al. Body condition varied annually, with years of significantly better and worse values. The body condition of lactating females was significantly worse than that of other whales at all times and was most often determined to be compromised.

Although there was no significant trend in scars over time, the annual percentage of animals observed with rope still on the body i. Parallel studies on entanglement scars in humpback whales suggest that entanglement is a major conservation issue in other species as well; the majority of humpback whales in the Gulf of Maine and also in northern Southeast Alaska suffer an entanglement at some point in their lives Neilson et al.

Entanglement has been traditionally viewed as an issue of immediate mortality, but an episode of entanglement may also induce long-term physiological costs even if the whale frees itself from the fishing gear, including reduced reproduction, reduced feeding, increased drag with swimming, and increased susceptibility to disease Knowlton and Kraus, ; Kot et al. For cetaceans, IR thermography can only assess the temperature of parts of the animal visible above the water surface, yet even this small portion of the animal can reveal interesting patterns in physiological responses to environmental changes and human stressors.

In dolphins, thermal imaging of the dorsal fin has been used to study effects of human disturbance e. Infrared studies in large whales date back to , when Cuyler et al. Infrared studies of large whales since then have primarily focused on detection of whale presence e. Overall, quantitative scoring of visual appearance has proved useful for monitoring nutritional status and general health. Visual appearance in several species has been shown to correlate directly to both reproduction and presumed mortality.

Challenges include the following: Aerial imaging provides a desirable whole-body perspective, but has the substantial drawbacks of cost and safety issues. However, it may be possible to use remote-controlled hexacopters as a low-cost option for aerial photography that could be operated from small boats at sea W. Another limitation of the aerial method is the difficulty of accurately assessing the maximal width at each point on the animal when it may be submerged in water that is variably opaque.

Next steps should amalgamate assessment of body condition in photo-identified whale populations with a better understanding of cumulative human impacts upon their rates of morbidity and mortality. For example, long-term photographic assessment of entanglement rates e. Such multi-analytical approaches would be very informative about the sublethal costs of human impacts.

In contrast to the dearth of physiological information available from large whales in the past, we now have three types of physiological samples that are fairly readily obtainable from whales faeces, blow, and biopsy samples as well as an increasing amount of physiological data that can be gleaned from visual assessment. All four methods are feasible to implement from a field perspective.

Many, if not most, cetacean research teams already routinely take photographs and collect biopsy samples during population surveys; many teams also have considerable skill with using cantilevered poles around whales for disentanglement, tagging, etc. Endocrine studies can potentially take advantage of three possible sample types—faeces, blow, and blubber—with all three matrices being likely to represent different time frames for acute vs. Furthermore, each sample type can provide separate information on specific organ systems, e. Finally, the possibility of developing a remote blood-sampling apparatus should not be discounted; creative engineering approaches might yet devise a solution to this difficult problem.

Whatever the sample type, the value of beginning with well-studied populations of known individuals cannot be overstated. For all the techniques discussed here, assay validation and careful development using populations with known individuals has been extremely useful. Specific examples include the well-studied NARW population in the Bay of Fundy, the southern resident killer whale population of Puget Sound, and certain populations of humpbacks. We encourage researchers to take advantage of well-known, photo-identified populations when translating novel analytical methods from terrestrial species and small captive odontocetes to large free-swimming whales.

When feasible, testing and validating methods with display or rehabilitated animals would also be invaluable.

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Once these techniques have been validated and proven in several baleen species and several odontocetes, we consider it likely that the techniques can then be applied to other populations that may not have known individuals, especially if study designs take advantage of comparison of populations before vs. We expect that combinations of different techniques will provide useful cross-checks and cross-validations.

For example, stress assessment, a common issue in conservation management, can be approached via multiple independent techniques, potentially including the following: Together, faecal samples, blow samples, biopsy samples, and visual assessment methods have the potential to revolutionize our understanding of large whale reproductive cycles, stress physiology, nutritional status, host immune response, pathogen and parasite load, and more.

Ultimately, these data can be used to assess how these physiological parameters are affected by the numerous conservation pressures impacting many large whale populations today. Further development of these techniques could identify which measures will be most useful as early warning indicators of potentially serious sublethal or even lethal impacts and which may indicate long-term chronic impacts.

As a next step, conservation physiology studies of large whales will benefit greatly from cross-disciplinary approaches that include cetacean conservation researchers along with experts from other fields e. We wish to thank the members of the North Atlantic Right Whale Consortium for collecting samples for right whale studies, permitting use of sightings and life history data, and for decades of dedicated research on these endangered whales.

We are grateful to Heather Pettis for assistance with compilation of the visual health-assessment figures, and Jodie Treloar for manuscript assistance. The content of this work is solely the responsibility of the authors and does not necessarily represent the official view of these agencies. National Center for Biotechnology Information , U. Journal List Conserv Physiol v. Published online May Moore , 2 Rosalind M. Rolland , 1 Nicholas M. Kellar , 3 Ailsa J. Hall , 4 Joanna Kershaw , 4 Stephen A.

Raverty , 5 Cristina E. Davis , 6 Laura C. Yeates , 7 Deborah A. Fauquier , 8 Teresa K. Rowles , 8 and Scott D. Hunt 1 John H. Rolland 1 John H. Kraus 9 John H. Author information Article notes Copyright and License information Disclaimer. This article has been cited by other articles in PMC. Abstract Large whales are subjected to a variety of conservation pressures that could be better monitored and managed if physiological information could be gathered readily from free-swimming whales.

Blow, biopsy dart, Cetacea, faecal samples, non-invasive, visual health assessment. Faecal samples Faecal samples have long been studied for information specific to digestive physiology, particularly intestinal parasitology and diet analysis, but more recently mammalian faeces have been shown to contain very high concentrations of steroid hormones as well as several other measures of interest Wasser et al.

Faecal sample collection from large whales Faecal samples have been collected successfully from multiple species of large whales, including North Atlantic right whales NARWs; Eubalaena glacialis ; Weisbrod et al. Open in a separate window. Faecal hormone analysis Many recent faecal studies in large whales have focused on faecal steroid hormones e. Non-endocrine faecal measures Faecal samples have long been used for diet analysis, traditionally via visual inspection of skeletal elements and more recently using genetic methods and stable isotope analysis e.

Targeted biomarker sampling possible May contain large variety of other detectable compounds? Respiratory sample collection from large whales Several of the human breath-sampling methods described above have been successfully modified for cetaceans. Blow hormones In , detectable testosterone and progesterone were reported in blow droplets collected from both humpbacks and NARWs, as assessed with liquid chromatography—mass spectrometry Hogg et al.

Blow microbiology Blow sampling may provide a unique window into the microbiology and, potentially, the immune status of large whales. Blow transcriptomics and gene expression Transcriptomics is the study of all the RNA produced by a cell, tissue, or entire organism at any given time, reflecting the genes that are being expressed at that point in time. Biopsy samples blubber and skin Since the early s, dart biopsying has become one of the most common collection methods for obtaining biological tissue samples from free-ranging cetaceans Lambertsen, ; Mathews et al. Biopsy sample collection methods for large whales Dart biopsy samples are predominantly collected using either a crossbow or a pneumatic rifle with modified dart tips Lambertsen, ; Mathews et al.

Blubber hormones Recently, a nascent but growing field of research has focused on measurement of lipophilic hormones, especially steroids, in blubber Mansour et al. Blubber contaminants Blubber biopsies have long been used to measure exposure to organic pollutants.

Blubber lipid analysis Blubber contains a variety of different fatty acids FAs , and the types and proportions of FAs present can reveal information relevant to dietary physiology, thermal physiology, and even the age of the individual. Epidermal diseases and microbiology Epidermal diseases in free-ranging whales and dolphins have been studied primarily using photographic analyses see next main section , but can also be studied via epidermal and blubber samples.

Epidermal proteomics A unique aspect of biopsy samples, in comparison to other matrices faeces and blow , is that biopsy samples contain large quantities of intact peptides and proteins. Epidermal transcriptomics and gene expression Transcriptomics has become a widely used approach for the study of normal and diseased gene expression in human skin Cole et al.

External appearance of the animal Visual assessment of the external appearance of an animal is a time-honoured technique for assessing the health and nutritional status of individuals. Imaging techniques for large whales Most imaging techniques to assess appearance have been based on photographs from planes Fig.

Aerial photography Initial aerial measurement of whales at sea primarily focused on measuring body length Best and Ruther, ; Angliss et al. Boat-based photography Lateral body condition assessment from a boat is only semi-quantitative, given the inability to observe the full submerged body outline, but data indicate that viewing even a portion of the animal may provide useful information on individual health. Summary and conclusions In contrast to the dearth of physiological information available from large whales in the past, we now have three types of physiological samples that are fairly readily obtainable from whales faeces, blow, and biopsy samples as well as an increasing amount of physiological data that can be gleaned from visual assessment.

Expert Rev Mol Diagn Mar Mamm Sci 8: Mar Mamm Sci Report of the International Whaling Commission , J Invest Dermatol J Exp Biol Sci Total Environ J Breath Res 3: J Breath Res 2: J Anal Toxicol Best PB, Ruther H. How far are we from clinical practice? Trends Analyt Chem Mar Pollut Bull Brewis IA, Brennan P. Adv Protein Chem Struct Biol Int J Antimicrob Agents Int Zoo Yearb Sci Total Environ — J Exp Mar Bio Ecol Curr Opin Pulm Med Anal Bioanal Chem Clapham PJ, Mattila D.

Mar Mamm Sci 9: Wound Repair Regen 9: Gen Comp Endocrinol Davidsson A, Schmekel B. J Clin Lab Anal Int J Pharm Annu Rev Mar Sci 4: J Breath Res 5: Dev Comp Immunol Mar Environ Res Mar Environ Res 69 Supplement 1: Endang Species Res 6: A novel method for DNA collection from cetacean blow. Int J Cancer Ganz T, Lehrer R.

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