Search

Odontocete cetaceans have undergone profound modifications to their integument and sensory systems and are generally thought to lack specialized exocrine glands that in terrestrial mammals function to produce chemical signals (Thewissen & Nummela, 2008). Keenan-Bateman et al. (2016, 2018), though, introduced an enigmatic exocrine gland, associated with the false gill slit pigmentation pattern in Kogia breviceps. These authors provided a preliminary description of this cervical gill slit gland in their helminthological studies of the parasitic nematode, Crassicauda magna. This study offers the first detailed gross and histological description of this gland and reports upon key differences between immature and mature individuals. Investigation reveals it is a complex, compound tubuloalveolar gland with a well-defined duct that leads to a large, and expandable central chamber, which in turn leads to two caudally projecting diverticula. All regions of the gland contain branched tubular and alveolar secretory regions, although most are found in the caudal diverticula, where the secretory process is holocrine. The gland lies between slips of cutaneous muscle, and is innervated by lamellar corpuscles, resembling Pacinian's corpuscles, suggesting that its secretory product may be actively expressed into the environment. Mature K. breviceps display larger gland size, and increased functional activity in glandular tissues, as compared to immature individuals. These results demonstrate that the cervical gill slit gland of K. breviceps shares morphological features of the specialized, chemical signaling, exocrine glands of terrestrial members of the Cetartiodactyla.

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.

Compared with terrestrial mammals, marine mammals possess increased muscle myoglobin concentrations (Mb concentration, g Mb · 100g−1 muscle), enhancing their onboard oxygen (O2) stores and their aerobic dive limit. Although myoglobin is not homogeneously distributed, cetacean muscle O2 stores have been often determined by measuring Mb concentration from a single muscle sample (longissimus dorsi) and multiplying that value by the animal’s locomotor muscle or total muscle mass. This study serves to determine the accuracy of previous cetacean muscle O2 stores calculations. For that, body muscles from three delphinid species: Delphinus delphis, Stenella coeruleoalba, and Stenella frontalis, were dissected and weighed. Mb concentration was calculated from six muscles/muscle groups (epaxial, hypaxial and rectus abdominis, mastohumeralis, sternohyoideus, and dorsal scalenus), each representative of different functional groups (locomotion powering swimming, pectoral fin movement, feeding and respiration, respectively). Results demonstrated that the Mb concentration was heterogeneously distributed, being significantly higher in locomotor muscles. Locomotor muscles were the major contributors to total muscle O2 stores (mean 92.8%) due to their high Mb concentration and large muscle masses. Compared to this method, previous studies assuming homogenous Mb concentration distribution likely underestimated total muscle O2 stores by 10% when only considering locomotor muscles and overestimated them by 13% when total muscle mass was considered.

Understanding toothed whale (odontocete) diving gas dynamics is important given the recent atypical mass strandings of odontocetes (particularly beaked whales) associated with mid-frequency naval sonar. Some stranded whales have exhibited gas emboli (pathologies resembling decompression sickness) in their specialized intramandibular and extramandibular fat bodies used for echolocation and hearing. These tissues have phylogenetically unique, endogenous lipid profiles with poorly understood biochemical properties. Current diving gas dynamics models assume an Ostwald nitrogen (N2) solubility of 0.07 ml N2 ml−1 oil in odontocete fats, although solubility in blubber from many odontocetes exceeds this value. The present study examined N2 solubility in the blubber and mandibular fats of seven species across five families, relating it to lipid composition. Across all species, N2 solubility increased with wax ester content and was generally higher in mandibular fats (0.083±0.002 ml N2 ml−1 oil) than in blubber (0.069±0.007 ml N2 ml−1 oil). This effect was more pronounced in mandibular fats with higher concentrations of shorter, branched fatty acids/alcohols. Mandibular fats of short-finned pilot whales, Atlantic spotted dolphins and Mesoplodon beaked whales had the highest N2 solubility values (0.097±0.005, 0.081±0.007 and 0.080±0.003 ml N2 ml−1 oil, respectively). Pilot and beaked whales may experience high N2 loads during their relatively deeper dives, although more information is needed about in vivo blood circulation to mandibular fats. Future diving models should incorporate empirically measured N2 solubility of odontocete mandibular fats to better understand N2 dynamics and potential pathologies from gas/fat embolism.

Growth in common bottlenose dolphins (Tursiops truncatus) was investigated through examination of sex-specific, ontogenetic changes in the mass of 38 discrete body compartments, utilizing stranded dolphins in good body condition (n = 145). Ontogenetic allometry and the body composition technique were used to quantitatively describe growth patterns. Although adult males were significantly larger than adult females in total body mass (TBM) and total length, overall patterns of growth were remarkably similar between sexes. The integument, locomotor muscle, and vertebral column together represented 50%–58% of TBM across all life history categories, although their relative contributions varied ontogenetically. Young dolphins invested the greatest percentage of TBM in integument, while locomotor muscle was the single largest body component in adults. In both sexes (1) most muscle groups displayed positive allometry, (2) most skeletal elements displayed negative allometric or isometric growth, (3) most abdominal viscera associated with digestion displayed positive allometry, and (4) the brain displayed negative allometric growth. Reproductive tissues exhibited the highest rates of growth in both sexes, and increased as a percentage of TBM with maturity. This study provides an integrated view of bottlenose dolphin growth and a quantitative baseline of body composition for future monitoring of this sentinel species of ecosystem health.

Odontocete echolocation clicks are generated by pneumatically driven phonic lips within the nasal passage, and propagated through specialized structures within the forehead. This study investigated the highly derived echolocation structures of the pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales through careful dissections (N = 18 K. breviceps, 6 K. sima) and histological examinations (N = 5 K. breviceps). This study is the first to show that the entire kogiid sound production and transmission pathway is acted upon by complex facial muscles (likely derivations of the m. maxillonasolabialis). Muscles appear capable of tensing and separating the solitary pair of phonic lips, which would control echolocation click frequencies. The phonic lips are enveloped by the “vocal cap,” a morphologically complex, connective tissue structure unique to kogiids. Extensive facial muscles appear to control the position of this structure and its spatial relationship to the phonic lips. The vocal cap's numerous air crypts suggest that it may reflect sounds. Muscles encircling the connective tissue case that surrounds the spermaceti organ may change its shape and/or internal pressure. These actions may influence the acoustic energy transmitted from the phonic lips, through this lipid body, to the melon. Facial and rostral muscles act upon the length of the melon, suggesting that the sound “beam” can be focused as it travels through the melon and into the environment. This study suggests that the kogiid echolocation system is highly tunable. Future acoustic studies are required to test these hypotheses and gain further insight into the kogiid echolocation system. Anat Rec, 298:1301–1326, 2015. © 2015 Wiley Periodicals, Inc.

Cetaceans span a large range of body sizes and include species with the largest known locomotor muscles. To date, force output and thrust production have only been directly measured in the common bottlenose dolphin (Tursiops truncatus), although thrust forces have been hydrodynamically modeled for some whales. In this study, two metrics of epaxial muscle size—cross-sectional area (CSA) and mass—were used to estimate force output for 22 species (n = 83 specimens) ranging in size from bottlenose dolphins to blue whales (Balaenoptera musculus). Relative to total body length (TL), maximum force output estimated based upon both muscle CSA (TL1.56 ± 0.05) and mass (TL2.64 ± 0.07) scaled at rates lower than those predicted by geometric scaling, suggesting relative force output decreases with increasing body size in cetaceans. Estimated maximal force outputs were compared to both published drag forces and to the breaking strengths of commercial fishing lines known to entangle whales. The breaking strengths of these lines are within the same order of magnitude, and in some cases, exceed the estimated maximal force output of whales. These results suggest that while powerful animals, large whales may be unable to break the extremely strong fishing line used today.

Electronic tags have proven to be valuable tools in assessing small cetacean movement and behavior. However, problems associated with tag size and attachment have limited duration and damaged dorsal fins. These outcomes have motivated researchers to develop a new satellite-linked tag design that reduces detrimental effects to tagged animals, while increasing transmission durations. The goals of this study were to review previous studies that deployed single-pin transmitters and determine factors that influence transmission duration. Then, test these factors utilizing computational fluid dynamics (CFD) models to identify an optimal single-pin satellite-linked tag design, and evaluate this prototype through field studies. A review of four projects, which deployed 77 single-pin radio tags, determined that tags attached along the lower third of the dorsal fin and approximately 33 mm from the trailing edge resulted in longer transmission durations and reduced negative impacts to the dorsal fin. Based upon these results and CFD modeling, prototype, single-pin satellite-linked tags (n = 25) transmitted for 163 ± 22 d (mean ± 95% CI) which greatly exceeded transmissions for previous small cetacean telemetry studies. These results suggest that the newly developed single-pin satellite-linked tag design strikes a balance between reducing impacts to the individual while maximizing transmissions.

When a marine mammal dives, breathing and locomotion are mechanically uncoupled, and its loco- motor muscle must power swimming when oxygen is lim- ited. The morphology of that muscle provides insight into both its oxygen storage capacity and its rate of oxygen consumption. This study investigated the m. longissimus dorsi, an epaxial swimming muscle, in the long duration, deep-diving pygmy sperm whale (Kogia breviceps) and the short duration, shallow-diving Atlantic bottlenose dol- phin (Tursiops truncatus). Muscle myoglobin content, fiber type profile (based upon myosin ATPase and succi- nate dehydrogenase assays), and fiber size were meas- ured for five adult specimens of each species. In addition, a photometric analysis of sections stained for succinate dehydrogenase was used to create an index of mitochon- drial density. The m. longissimus dorsi of K. breviceps dis- played significantly a) higher myoglobin content, b) larger proportion of Type I (slow oxidative) fibers by area, c) larger mean fiber diameters, and d) lower indices of mi- tochondrial density than that of T. truncatus. Thus, this primary swimming muscle of K. breviceps has greater ox- ygen storage capacity, reduced ATP demand, and likely a reduced rate of oxygen consumption relative to that of T. truncatus. The locomotor muscle of K. breviceps appears able to ration its high onboard oxygen stores, a feature that may allow this species to conduct relatively long du- ration, deep dives aerobically. J. Morphol. 274:663-675, 2013. ! 2013 Wiley Periodicals, Inc.

Blubber, a specialized form of subdermal adipose tissue, surrounds marine mammal bodies. Typically, adipose tissue is perfused by capillaries but information on blubber vascularization is lacking. This study's goals were to: 1) describe and compare the microvasculature (capillaries, microarterioles, and microvenules) of blubber across odontocete species; 2) compare microvasculature of blubber to adipose tissue; and 3) examine relationships between blubber's lipid composition and its microvasculature. Percent microvascularity, distribution, branching pattern, and diameter of microvessels were determined from images of histochemically stained blubber sections from shallow-diving bottlenose dolphins (Tursiops truncatus), deeper-diving pygmy sperm whales (Kogia breviceps), deep-diving beaked whales (Mesoplodon densirostris; Ziphius cavirostris), and the subdermal adipose tissue of domestic pigs (Sus scrofa). Tursiops blubber showed significant stratification in percent microvascularity among the superficial, middle, and deep layers and had a significantly higher percent microvascularity than all other animals analyzed, in which the microvasculature was more uniformly distributed. The percent microvasculature of Kogia blubber was lower than that of Tursiops but higher than that of beaked whales and the subdermal adipose tissue of domestic pigs. Tursiops had the most microvascular branching. Microvessel diameter was relatively uniform in all species. There were no clear patterns associating microvascular and lipid characteristics. The microvascular characteristics of the superficial layer of blubber resembled the adipose tissue of terrestrial mammals, suggesting some conservation of microvascular patterns in mammalian adipose tissue. The middle and deep layers of blubber, particularly in Tursiops, showed the greatest departure from typical mammalian microvascular arrangement. Factors such as metabolics or thermoregulation may be influencing the microvasculature in these layers. © J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.

SUMMARYIn Atlantic bottlenose dolphins Tursiops truncatus, both the thickness and lipid content of blubber vary across ontogeny and across individuals of differing reproductive and nutritional status. This study investigates how these changes in blubber morphology and composition influence its thermal properties. Thermal conductivity (W m–1deg.–1, where deg. is °C) and thermal insulation(m2 deg. W–1) of dolphin blubber were measured in individuals across an ontogenetic series (fetus through adult, N=36),pregnant females (N=4) and emaciated animals (N=5). These thermal properties were determined by the simultaneous use of two common experimental approaches, the heat flux disc method and the standard material method. Thickness, lipid and water content were measured for each blubber sample. Thermal conductivity and insulation varied significantly across ontogeny. Blubber from fetuses through sub-adults was less conductive(range=0.11–0.13±0.02 W m–1deg.–1) than that of adults (mean=0.18 W m–1deg.–1). The conductivity of blubber from pregnant females was similar to non-adult categories, while that of emaciated animals was significantly higher (0.24 ± 0.04 W m deg.–1) than all other categories. Blubber from sub-adults and pregnant females had the highest insulation values while fetuses and emaciated animals had the lowest. In nutritionally dependant life history categories, changes in blubber's thermal insulation were characterized by stable blubber quality (i.e. conductivity)and increased blubber quantity (i.e. thickness). In nutritionally independent animals, blubber quantity remained stable while blubber quality varied. A final, unexpected observation was that heat flux measurements at the deep blubber surface were significantly higher than that at the superficial surface, a pattern not observed in control materials. This apparent ability to absorb heat, coupled with blubber's fatty acid composition, suggest that dolphin integument may function as a phase change material.

Cetacean brains are among the least studied mammalian brains because of the formidable histological preparations of such relatively rare and large specimens. Although the bottlenose dolphin, Tursiops truncatus, has been the most extensively studied cetacean species, there have been relatively few studies of the brain of the infant bottlenose dolphin. In this study, we present the first magnetic resonance imaging (MRI)-based study of the brain of an infant bottlenose dolphin. Magnetic resonance images in the coronal plane were originally acquired and used to digitally generate a set of resectioned virtual images in orthogonal planes. A sequential set of images in all three planes was anatomically labeled and reveals major neuroanatomical features. Some of the distinctive features of cetacean brains are already evident in the infant bottlenose dolphin brain, while other features may represent differences that deserve further study.

This study investigated the functional morphology of the blubber that forms the caudal keels of the harbor porpoise (Phocoena phocoena). Blubber is a pliant biocomposite formed by adipocytes and structural fibers composed of collagen and elastic fibers. Caudal keels are dorsally and ventrally placed triangular wedges of blubber that define the hydrodynamic profile of the porpoise tailstock. Mechanical tests on carcasses demonstrate that when keels are bent, they strain nonuniformly along their lengths, with highest strains just caudal to the dorsal fin and lowest at the insertion of the flukes. Therefore, caudal keels undergo nonuniform longitudinal deformation while maintaining a stable, triangular cross-sectional shape. Polarizing and transmitted light microscopy techniques were used to investigate blubber's 3D fiber architecture along the length of the dorsal keel. The triangular cross-sectional shape of the keel appears to be maintained by structural fibers oriented to act as tensile stays. The construction of the blubber composite is regionally specific :structural fiber densities and diameters are higher in the relatively stiff caudal region of the keel than in the more deformable cranial keel region. The orientations of structural fibers also change along the length of the keel. Cranially, no fibers are oriented along the long axis, whereas a novel population of longitudinally oriented fibers reinforces the keel at the insertion of the flukes. Thus, differences in the distribution and orientation of structural fibers contribute to the regionally specific mechanical properties of the dorsal keel.

Nine bottlenose dolphin (Tursiops truncatus) calves that stranded in Virginia in 1996 and 1997 died of severe blunt-force trauma. Injuries were concentrated on the head and chest and multiple rib fractures, lung lacerations, and soft tissue contusions were prominent. Skeletal and/or soft tissue trauma occurred bilaterally in all of the calves. One had a bite wound across the left mandible that exhibited deep punctures consistent with the tooth placement in an adult bottlenose dolphin. The lesions were not compatible with predation, boat strike, fisheries interactions, rough-surf injury, or blast injury. However, they were similar to traumatic injuries described in stranded bottlenose dolphin calves and harbor porpoises (Phocoena phocoena) in Great Britain attributed to violent dolphin interactions. The evidence suggests that violent dolphin behavior was the cause of the trauma in the nine calves reported here and that infanticide occurs in bottlenose dolphins of the western North Atlantic.

The reproductive organs of Florida manatees (Trichechus manatus latirostris) are surrounded by thermogenic locomotory muscles and insulating fat. Manatees are reported to maintain core body temperatures of 35.6°–36.4° C, temperatures known to interfere with production and maturation of viable sperm in terrestrial mammals. We describe two novel venous plexuses associated with the manatee epididymis. Each epididymis is located in a hypogastric fossa at the caudolateral extremity of the abdominal cavity. Each hypogastric fossa is lined by an inguinal venous plexus that receives cooled blood from a superficial thoracocaudal plexus. We conclude that male manatees may prevent hyperthermic insult to their reproductive tissues by feeding cooled superficial blood to venous plexuses deep within their bodies. Female manatees also possess hypogastric fossae and venous structures similar to those found in male manatees. The ovaries, uterine tubes, and distal tips of the uterine horns are located in the hypogastric fossae. We suggest that the thermovascular structures we describe also prevent hypothermic insult to female manatee reproductive tissues. The venous structures in manatees are functionally similar to structures associated with reproductive thermoregulation in cetaceans and phocid seals. Thus, these thermovascular structures appear to be convergent morphological adaptations that occur in three clades of diving mammals with independent evolutionary histories. Anat Rec 264:339–347, 2001. Published 2001 Wiley-Liss, Inc.

At birth, the locomotor muscles of precocial, terrestrial mammals are similar to those of adults in both mass, as a percent of total body mass, and fiber-type composition. It is hypothesized that bottlenose dolphins (Tursiops truncatus), marine mammals that swim from the instant of birth, will also exhibit precocial development of locomotor muscles. Body mass data from neonatal and adult dolphins are used to calculate Grand's (1992) Neural and Muscular Indices of Development. Using these indices, the bottlenose dolphin is a Condition "3.5" neonate, where Condition 4 is the documented extreme of precocial development in terrestrial mammals. Moreover, myosin ATPase (alkaline preincubation) analyses of the epaxial locomotor m. extensor caudae lateralis show that neonatal dolphins have fiber-type profiles very similar to those of adults. Thus, based on mass and myosin ATPase activity, muscle development in dolphins is precocial. However, succinic dehydrogenase and Nile red histochemistry demonstrate that neonatal dolphin muscle has mitochondrial and lipid distributions different from those found in adults. These data suggest that neonates have a lower aerobic capacity than adults. Dolphin neonates may compensate for an apparent lack of aerobic stamina in two ways: 1) by being positively buoyant, with a relatively increased investment of their total body mass in blubber, and 2) by "free-riding" off their mothers. This study investigates quantitatively the development of a dolphin locomotor muscle and offers suggestions about adaptations required for a completely aquatic existence.

The similarity in swimming style and external body shape between dolphins and scombrid fishes, especially tunas, is a textbook example of evolutionary convergence. I identify additional morphological features of the musculoskeletal system shared by dolphins and tunas. Specifically, these swimmers share a pattern of force transmission through a complex, three-dimensional system of collagenous fabrics, which are stiffened by muscular hydrostatic pressure. This force transmission system increases both the displacement advantage and moment arm of contracting axial muscle. These features represent a functionally significant design for steady swimming vertebrates.

The subdermal connective tissue sheath (SDS) of dolphins is a fibre-reinforced membrane connected to other locomotor tissues, including blubber, axial muscles and tendons, and vertebral column. The complicated connections between the SDS and other locomotor tissues suggest that the SDS acts as a peripheral skeletal element for the axial locomotor muscles and as an anchor for a de novo dermal appendage, the dorsal fin. The morphology of the SDS suggests that the dolphin can be modelled as a fibre-wound, thin-walled, pressurized cylinder. Existing cylinder models predict that the SDS functions to resist torsional forces, prevent aneurysms, and limit wrinkling when the dolphin bends in locomotion. I present a new functional model that more accurately represents the morphology of the dolphin cylinder wrapped by the SDS. The new model predicts that the SDS: (1) acts as a retinaculum for the terminal tendons of the axial locomotor muscles; and (2) plays a role in maintaining the laterally flattened cross-sectional shape of the caudal peduncle. The model is based on external morphological features of dolphins shared by other steady swimming aquatic vertebrates, such as carangiform and thunniform fishes. These features, which include a streamlined body shape and narrow necking of the caudal peduncle have been identified as adaptions to reduce drag. The new model offers insight into some of the structural features of the body wall required to maintain the hydrodynamicallytuned, external morphology of steady-swimming vertebrates.

The dolphin upstroke is powered primarily by the m. multifidus and m. longissimus—robust muscles that insert serially along the vertebral column. Intramuscular and tendon morphology coupled with kinematic data are used to hypothesize regionally specific functions of these muscles. Both muscles develop equivalent forces (approximately 2kN) in the region of the thoraco-lumbar spine, but transmit those forces to different regions of the vertebral column. The m. multifidus transmits the majority of its force locally to the thoraco-lumar spine, a region of the body that undergoes no measurable bending. The action of the m. multifidus appears to be to stiffen its deep tendon of insertion, forming a temporary skeletal element for the m. longissimus. The m. longissimus transmits the majority of its force to the caudal spine, by way of a novel interaction between its insertional tendons and the subdermal connective tissue sheath. This insertional pattern confers torsional stiffness on the caudal peduncle, and allows the m. longissimus to do relatively more work (118 J) than if it had a typical mammalian insertional pattern. The caudal extension of the m. multifidus does 12 J of work on the caudal spine during an upstroke. The caudal extension of the m. longissimus transmits its force to the vertebrae in the caudal flukes and is the only epaxial muscle that acts to control the angle of attack of the fluke blade.

In Atlantic bottlenose dolphins (Tursiops truncatus) the thickness and lipid content of blubber (the integument's specialized hypodermis) varies across ontogeny and with reproductive and nutritional state. Because the integument comprises up to 25% of total body mass in this species, ontogenetic changes in its lipid content may influence whole body buoyancy. The density and volume of the integument were measured and its buoyancy calculated across an ontogenetic series of dolphins and in pregnant and emaciated adults (total n = 45). Regional differences between the metabolically labile trunk integument and the structural tailstock integument were also investigated. Mean densities of both trunk and tailstock integument were similar across life history categories (trunk = 1,040.7 ± 14.1 kg/m 3 ; tailstock = 1,077.1 ± 21.2 kg/m 3 ) and were statistically similar to the density of seawater (1,026 kg/m 3 ). The mean buoyant force of integument from the trunk (—1.01 ± 1.74 N) and tailstock (—0.30 ± 0.21 N) did not vary significantly across ontogeny. In contrast, pregnancy and emaciation did influence the integument's buoyancy, which ranged between 9 N and —45 N in these categories. Although neutral during growth, the integument's contribution to whole body buoyancy can be influenced by an individual's reproductive and nutritional status.

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 appear to be coupled only during their brief surfacing event, when they respire explosively (up to 90% of total lung volume in approximately 0.3s) (Ridgway et al., 1969). The predominantly slow-twitch fiber profile of their diaphragm (Dearolf, 2003) suggests that this muscle does not likely power their rapid ventilatory event. Based upon Bramble’s (1989) biomechanical model of locomotor-respiratory coupling in galloping mammals, I hypothesized that muscles located within the cranial-cervical and lumbo-pelvic units, which act upon the thoracic unit, function to power ventilation in bottlenose dolphins. I also hypothesized that these muscles would be composed predominantly of fast-twitch fibers to facilitate the bottlenose dolphin’s rapid ventilation. The gross morphology (n=6) of cranio-cervical (sternomastoid, sternohyoid, scalenes), thoracic (intercostals), and lumbo-pelvic (rectus abdominis, abdominal obliques, hypaxialis) muscles and the fiber-type profiles (n=6) of selected muscles (sternohyoid, sternomastoid, and 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 cranio-cervical muscles act to draw the sternum and associated ribs cranio-dorsally, which flares the ribs laterally, and increases thoracic cavity volume required for inspiration. The thoracic muscles physically link the ribs to create a single functional unit; these muscles can also act to control the size of the intercostal space. The lumbo-pelvic muscles act to draw the sternum and caudal ribs caudally, which decreases the volume of the thoracic and abdominal cavities required for expiration. All muscles investigated were composed predominantly of fast-twitch fibers (range 72-88% 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. However, the mechanisms that permit dolphins to selectively couple and uncouple their locomotor and ventilatory systems, depending upon whether they are respiring at the surface or swimming on a breath-hold, remain unknown.

Foetal dolphins (Tursiops truncatus) are bent ventrolaterally, such that the tailflukes and lower jaw are juxtaposed. The lateral flexibility required en utero may compromise the efficiency of the dorsoventral oscillations required of the swimming neonate. The m. intertransversarius caudae dorsalis (IT) is the most laterally placed epaxial muscle. Bilateral contractions of the IT could limit lateral deformations of the flexible tailstock of the early neonate. We test the hypothesis that the IT is functioning as a postural muscle in neonates by examining its morphological, histological and biochemical properties. The neonatal IT has a relatively large cross-sectional area and bending moment, as well as a large proportion of slow-twitch fibres and elevated myoglobin concentrations. Our results demonstrate that the IT is functionally capable of performing this specific postural function in neonatal dolphins. In later life-history stages, when postural control is no longer needed, the IT serves to fine-tune the position of the tailstock during locomotion. The changing function of the adult IT is concomitant with changes in morphology and biochemistry, and most notably, with an increase in the proportion of fast-twitch fibres. We suggest that these changes reflect strong selective pressure to improve locomotor abilities by limiting lateral deformations during this critical life-history stage.

SUMMARYThe dorsal fin of the bottlenose dolphin Tursiops truncatuscontains blood vessels that function either to conserve or to dissipate body heat. Prior studies have demonstrated that heat flux, measured from a single position on the dorsal fin, decreases during body cooling and diving bradycardia and increases after exercise and at the termination of the dive response. While prior studies attributed changes in heat flux to changes in the pattern of blood flow, none directly investigated the influence of vascular structures on heat flux across the dorsal fin. In this study we examined whether heat flux is higher directly over a superficial vein,compared to a position away from a vein, and investigated the temporal relationship between heart rate, respiration and heat flux. Simultaneous records of heat flux and skin temperature at three positions on the dorsal fins of 19 wild bottlenose dolphins (with the fin in air and submerged) were collected, together with heart rate and respiration. When the fin was submerged, heat flux values were highest over superficial veins, usually at the distal tip, suggesting convective delivery of heat, via blood, to the skin's surface. Conversely, in air there was no relationship between heat flux and superficial vasculature. The mean difference in heat flux (48 W m-2) measured between the three fin positions was often equal to or greater than the heat flux that had been recorded from a single position after exercising and diving in prior studies. Tachycardia at a respiratory event was not temporally related to an increase in heat flux across the dorsal fin. This study suggests that the dorsal fin is a spatially heterogeneous thermal surface and that patterns of heat flux are strongly influenced by underlying vasculature.

Cetacean (dolphin, whale, and porpoise) brains are among the least studied mammalian brains because of the formidable challenge of collecting and histologically preparing such relatively rare and large specimens. Magnetic resonance imaging offers a means of observing the internal structure of the brain when traditional histological procedures are not practical. Furthermore, internal structures can be analyzed in their precise anatomic positions, which is difficult to accomplish after the spatial distortions often accompanying histological processing. In this study, images of the brain of an adult bottlenose dolphin, Tursiops truncatus, were scanned in the coronal plane at 148 antero-posterior levels. From these scans a computer-generated three-dimensional model was constructed using the programs VoxelView and VoxelMath (Vital Images, Inc.). This model, wherein details of internal and external morphology are represented in three-dimensional space, was then resectioned in orthogonal planes to produce corresponding series of virtual sections in the horizontal and sagittal planes. Sections in all three planes display the sizes and positions of major neuroanatomical features such as the arrangement of cortical lobes and subcortical structures such as the inferior and superior colliculi, and demonstrate the utility of MRI for neuroanatomical investigations of dolphin brains.

The cetacean reproductive system is surrounded by thermogenic locomotory muscle and insulating blubber. This arrangement suggests elevated temperatures at the uterus that could induce detrimental effects on fetal development. We present anatomical evidence for a complex countercurrent heat exchange system that could function to regulate the thermal environment of the uterus and a developing fetus. Cooled venous blood from the surfaces of the dorsal fin and flukes enters the abdominal cavity via the lumbo-caudal venous plexus. This plexus is juxtaposed to the arterial and venous plexuses associated with the uterus. The morphology of the lumbo-caudal venous plexus suggests that it acts as a "heat sink" for the adjacent tissues. Heat may be transferred to the cool, lumbo-caudal venous plexus from the warm blood in the arterial and venous plexuses supplying the uterus. Heat may also be transferred from adjacent locomotory muscles to the cool lumbo-caudal venous plexus. The countercurrent heat exchanger created by the juxtaposition of the lumbo-caudal venous plexus with the uterovarian arterial plexus is similar in design to that of the countercurrent heat exchanger described for male cetaceans. The functional implications of introducing cool superficial blood into the abdominal cavity of a diving, and locomoting female cetacean are discussed.

I do not think that anyone who has not dissected the spinal muscles of a cetacean can quite appreciate the impracticability of homologizing exactly all of the elements.