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The cetacean vertebral canal houses the spinal cord and arterial supply to and venous drainage from the entire central nervous system (CNS). Thus, unlike terrestrial mammals, the cetacean spinal cord lies within a highly vascularized space. We compared spinal cord size and vascular volumes within the vertebral canal across a sample of shallow and deep diving odontocetes. We predicted that the (a) spinal cord, a metabolically expensive tissue, would be relatively small, while (b) volumes of vascular structures would be relatively large, in deep versus shallow divers. Our sample included the shallow diving Tursiops truncatus (n = 2) and Delphinus delphis (n = 3), and deep diving Kogia breviceps (n = 2), Mesoplodon europaeus (n = 2), and Ziphius cavirostris (n = 1). Whole, frozen vertebral columns were cross-sectioned at each intervertebral disc, scaled photographs of vertebral canal contents acquired, and cross-sectional areas of structures digitally measured. Areas were multiplied by vertebral body lengths and summed to calculated volumes of neural and vascular structures. Allometric analyses revealed that the spinal cord scaled with negative allometry (b = 0.51 ± 0.13) with total body mass (TBM), and at a rate significantly lower than that of terrestrial mammals. As predicted, the spinal cord represented a smaller percentage of the total vertebral canal volume in the deep divers relative to shallow divers studied, as low as 2.8% in Z. cavirostris. Vascular volume scaled with positive allometry (b = 1.2 ± 0.22) with TBM and represented up to 96.1% (Z. cavirostris) of the total vertebral canal volume. The extreme deep diving beaked whales possessed 22-35 times more vascular volume than spinal cord volume within the vertebral canal, compared with the 6-10 ratio in the shallow diving delphinids. These data offer new insights into morphological specializations of neural and vascular structures that may contribute to differential diving capabilities across odontocete cetaceans., (© 2021 Wiley Periodicals LLC.)

Beaked whales are a poorly known but diverse group of whales that have received considerable attention due to strandings that have been temporally and spatially associated with naval sonar deployment. Postmortem studies on stranded carcasses have revealed lesions consistent with decompression sickness, including intravascular gas and fat emboli. These findings have been supported by analyses of intravascular gas emboli showing composition dominated by nitrogen gas. To increase our understanding of the pathophysiology of nitrogen bubble formation and intravascular embolization, we examined the gross and microscopic anatomy of the venous system in the head of beaked whales. Since the potential sources of intravascular fat and gas emboli were of greatest interest, focus was placed on the acoustic fat bodies and pneumatic accessory sinus system. Herein, we describe intimate arteriovenous associations with specialized adipose depots and air sinuses in beaked whales. These vascular structures comprise an extensive network of thin-walled vessels with a large surface area, which is likely to facilitate exchange of nitrogen gas and may, therefore, form anatomic regions that may be important in physiological management of diving gases. These structures may also be vulnerable to pathologic introduction of emboli into the vascular system. Expansive, thin-walled venous lakes are found within the pterygoid region, which suggest the potential for nitrogen exchange as well as for compensation of middle-ear pressures during descent on a dive. These findings warrant further research into the structure and function of this morphology as it relates to normal and pathologic physiology., (© 2015 Wiley Periodicals, Inc.)

Beaked whales are medium-sized toothed whales that inhabit depths beyond the continental shelf; thus beaked whale strandings are relatively infrequent compared to those of other cetaceans. Beaked whales have been catapulted into the spotlight by their tendency to strand in association with naval sonar deployment. Studies have shown the presence of gas and fat emboli within the tissues and analysis of gas emboli is suggestive of nitrogen as the primary component. These findings are consistent with human decompression sickness (DCS) previously not thought possible in cetaceans. Because, tissue loading with nitrogen gas is paramount for the manifestation of DCS and nitrogen loading depends largely on the vascular perfusion of the tissues, we examined the anatomy of the extracranial arterial system using stranded carcasses of 16 beaked whales from five different species. Anatomic regions containing lipid and/or air spaces were prioritized as potential locations of nitrogen gas absorption due to the known solubility of nitrogen in adipose tissue and the nitrogen content of air, respectively. Attention was focused on the acoustic fat bodies and accessory sinus system on the ventral head. We found much of the arterial system of the head to contain arteries homologous to those found in domestic mammals. Robust arterial associations with lipid depots and air spaces occurred within the acoustic fat bodies of the lower jaw and pterygoid air sacs of the ventral head, respectively. Both regions contained extensive trabecular geometry with small arteries investing the trabeculae. Our findings suggest the presence of considerable surface area between the arterial system, and the intramandibular fat bodies and pterygoid air sacs. Our observations may provide support for the hypothesis that these structures play an important role in the exchange of nitrogen gas during diving., (© 2015 Wiley Periodicals, Inc.)

Shallow-diving, coastal bottlenose dolphins (Tursiops truncatus) and deep-diving, pelagic pygmy and dwarf sperm whales (Kogia breviceps and K. sima) will experience vastly different ambient pressures at depth, which will influence the volume of air within their lungs and potentially the degree of thoracic collapse they experience. This study tested the hypotheses that lung size will be reduced and/or thoracic mobility will be enhanced in deeper divers. Lung mass (T. truncatus, n = 106; kogiids, n = 18) and lung volume (T. truncatus, n = 5; kogiids, n = 4), relative to total body mass, were compared. One T. truncatus and one K. sima were cross-sectioned to calculate lung, thoracic vasculature, and other organ volumes. Excised thoraxes (T. truncatus, n = 3; kogiids, n = 4) were mechanically manipulated to compare changes in thoracic cavity shape and volume. Kogiid lungs were half the mass and one-fifth the volume of those of similarly sized T. truncatus. The lungs occupied only 15% of the total thoracic cavity volume in K. sima and 37% in T. truncatus. The kogiid and dolphin thoraxes underwent similar changes in shape and volume, although the width of the thoracic inlet was relatively constrained in kogiids. A broader phylogenetic comparison demonstrated that the ratio of lung mass to total body mass in kogiids, physeterids, and ziphiids was similar to that of terrestrial mammals, while delphinids and phocoenids possessed relatively large lungs. Thus, small lung size in deep-diving odontocetes may be a plesiomorphic character. The relatively large lung size of delphinids and phocoenids appears to be a derived condition that may permit the lung to function as a site of respiratory gas exchange throughout a dive in these rapid breathing, short-duration, shallow divers.

Most mammals possess stamina because their locomotor and respiratory (i.e., ventilatory) systems are mechanically coupled. These systems are decoupled, however, in bottlenose dolphins (Tursiops truncatus) as they swim on a breath hold. Locomotion and ventilation are coupled only during their brief surfacing event, when they respire explosively (up to 90% of total lung volume in approximately 0.3 s) (Ridgway et al. 1969 Science 166:1651-1654). The predominantly slow-twitch fiber profile of their diaphragm (Dearolf 2003 J Morphol 256:79-88) suggests that this muscle does not likely power their rapid ventilatory event. Based on Bramble's (1989 Amer Zool 29:171-186) biomechanical model of locomotor-respiratory coupling in galloping mammals, it was hypothesized that locomotor muscles function to power ventilation in bottlenose dolphins. It was further hypothesized that these muscles would be composed predominantly of fast-twitch fibers to facilitate the bottlenose dolphin's rapid ventilation. The gross morphology of craniocervical (scalenus, sternocephalicus, sternohyoid), thoracic (intercostals, transverse thoracis), and lumbopelvic (hypaxialis, rectus abdominis, abdominal obliques) muscles (n = 7) and the fiber-type profiles (n = 6) of selected muscles (scalenus, sternocephalicus, sternohyoid, rectus abdominis) of bottlenose dolphins were investigated. Physical manipulations of excised thoracic units were carried out to investigate potential actions of these muscles. Results suggest that the craniocervical muscles act to draw the sternum and associated ribs craniodorsally, which flares the ribs laterally, and increases the thoracic cavity volume required for inspiration. The lumbopelvic muscles act to draw the sternum and caudal ribs caudally, which decreases the volumes of the thoracic and abdominal cavities required for expiration. All muscles investigated were composed predominantly of fast-twitch fibers (range 61-88% by area) and appear histochemically poised for rapid contraction. These combined results suggest that dolphins utilize muscles, similar to those used by galloping mammals, to power their explosive ventilation., ((c) 2008 Wiley-Liss, Inc.)

The melon is a lipid-rich structure located in the forehead of odontocetes that functions to propagate echolocation sounds into the surrounding aquatic environment. To date, the melon's ability to guide and impedance match biosonar sounds to seawater has been attributed to its unique fatty acid composition. However, the melon is also acted upon by complex facial muscles derived from the m. maxillonasolabialis. The goal of this study was to investigate the gross morphology of the melon in bottlenose dolphins (Tursiops truncatus) and to describe how it is tendinously connected to these facial muscles. Standard gross dissection (N = 8 specimens) and serial sectioning (N = 3 specimens) techniques were used to describe the melon and to identify its connections to the surrounding muscles and blubber in three orthogonal body planes. The dolphin forehead was also thin-sectioned in three body planes (N = 3 specimens), and polarized light was used to reveal the birefringent collagen fibers within and surrounding the melon. This study identified distinct regions of the melon that vary in shape and display locally specific muscle-tendon morphologies. These regions include the bilaterally symmetric main body and cone and the asymmetric right and left caudal melon. This study is the first to identify that each caudal melon terminates in a lipid cup that envelopes the echolocation sound generators. Facial muscles of the melon have highly organized tendon populations that traverse the melon and insert into either the surrounding blubber, the connective tissue matrix of the nasal plug, or the connective tissue sheath surrounding the sound generators. The facial muscles and tendons also lie within multiple orthogonal body planes, which suggest that the melon is capable of complex shape change. The results of this study suggest that these muscles could function to change the frequency, beam width, and directionality of the emitted sound beam in bottlenose dolphins. The echolocation sound propagation pathway within the dolphin forehead appears to be a tunable system., (Copyright 2008 Wiley-Liss, Inc.)