Vertebral morphology in extant porpoises: Radiation and functional implications.

Vertebral morphology has profound biomechanical implications and plays an important role in adaptation to different habitats and foraging strategies for cetaceans. Extant porpoise species (Phocoenidae) display analogous evolutionary patterns in both hemispheres associated with convergent evolution to coastal versus oceanic environments. We employed 3D geometric morphometrics to study vertebral morphology in five porpoise species with contrasting habitats: the coastal Indo-Pacific finless porpoise (Neophocaena phocaenoides); the mostly coastal harbor porpoise (Phocoena phocoena) and Burmeister's porpoise (Phocoena spinipinnis); and the oceanic spectacled porpoise (Phocoena dioptrica) and Dall's porpoise (Phocoenoides dalli). We evaluated the radiation of vertebral morphology, both in size and shape, using multivariate statistics. We supplemented data with samples of an early-radiating delphinoid species, the narwhal (Monodon monoceros); and an early-radiating delphinid species, the white-beaked dolphin (Lagenorhynchus albirostris). Principal component analyses were used to map shape variation onto phylogenies, and phylogenetic constraints were investigated through permutation tests. We established links between vertebral morphology and movement patterns through biomechanical inferences from morphological presentations. We evidenced divergence in size between species with contrasting habitats, with coastal species tending to decrease in size from their estimated ancestral state, and oceanic species tending to increase in size. Regarding vertebral shape, coastal species had longer centra and shorter neural processes, but longer transverse processes, while oceanic species tended to have disk-shaped vertebrae with longer neural processes. Within Phocoenidae, the absence of phylogenetic constraints in vertebral morphology suggests a high level of evolutionary lability. Overall, our results are in accordance with the hypothesis of speciation within the family from a coastal ancestor, through adaptation to particular habitats. Variation in vertebral morphology in this group of small odontocetes highlights the importance of environmental complexity and particular selective pressures for the speciation process through the development of adaptations that minimize energetic costs during locomotion and prey capture.

Genital interactions during simulated copulation among marine mammals.

Genitalia are morphologically variable across many taxa and in physical contact during intromission, but little is known about how variation in form correlates with function during copulation. Marine mammals offer important insights into the evolutionary forces that act on genital morphology because they have diverse genitalia and are adapted to aquatic living and mating. Cetaceans have a fibroelastic penis and muscular vaginal folds, while pinnipeds have a baculum and lack vaginal folds. We examined copulatory fit in naturally deceased marine mammals to identify anatomical landmarks in contact during copulation and the potential depth of penile penetration into the vagina. Excised penises were artificially inflated to erection with pressurized saline and compared with silicone vaginal endocasts and within excised vaginas in simulated copulation using high-resolution, diffusible iodine-based, contrast-enhanced computed tomography. We found evidence suggestive of both congruent and antagonistic genital coevolution, depending on the species. We suggest that sexual selection influences morphological shape. This study improves our understanding of how mechanical interactions during copulation influence the shape of genitalia and affect fertility, and has broad applications to other taxa and species conservation.

Unique feeding morphology in a new prognathous extinct porpoise from the Pliocene of California.

Modern porpoises (Odontoceti: Phocoenidae) are some of the smallest cetaceans and usually feed near the seafloor on small fish and cephalopods [1-3]. Within both extinct and extant phocoenids, no evidence for specialized mandibular morphology has been documented [4-7]. Here we describe a new species of extinct porpoise, Semirostrum ceruttii, from the marine Pliocene San Diego (4.2-1.6 mega-annum, Ma) and Purisima (5-2.5 Ma) formations of California. The mandibles comprise a long, fused, and nearly edentulous prognathous symphysis, extending farther beyond the rostrum than in any known mammal. Phylogenetic analyses based on morphology reconstruct Semirostrum ceruttii as sister to extant (crown) porpoise species with moderate support. We describe the spectacularly preserved holotype specimen based on computed tomography (CT) scans, which allowed visualization of the elongate mental and accessory canals within the symphysis. The elongate canals are similar to those found in Rynchops birds [8] and were likely involved in sensory function. Oblique labial wear facets present on numerous small conical mandibular teeth posterior to the symphysis suggest regular contact with benthic substrate. The unique mandibular and dental characteristics, along with robust scapulae, sternum, and unfused cervical vertebrae, support the interpretation that this species employed a form of benthic skim feeding by using its mandible to probe for and obtain prey.

A comparative analysis of marine mammal tracheas.

In 1940, Scholander suggested that stiffened upper airways remained open and received air from highly compressible alveoli during marine mammal diving. There are few data available on the structural and functional adaptations of the marine mammal respiratory system. The aim of this research was to investigate the anatomical (gross) and structural (compliance) characteristics of excised marine mammal tracheas. Here, we defined different types of tracheal structures, categorizing pinniped tracheas by varying degrees of continuity of cartilage (categories 1-4) and cetacean tracheas by varying compliance values (categories 5A and 5B). Some tracheas fell into more than one category along their length; for example, the harbor seal (Phoca vitulina) demonstrated complete rings cranially, and as the trachea progressed caudally, tracheal rings changed morphology. Dolphins and porpoises had less stiff, more compliant spiraling rings while beaked whales had very stiff, less compliant spiraling rings. The pressure-volume (P-V) relationships of isolated tracheas from different species were measured to assess structural differences between species. These findings lend evidence for pressure-induced collapse and re-inflation of lungs, perhaps influencing variability in dive depth or ventilation rates of the species investigated.

Comparative morphology of porpoise (Cetacea: Phocoenidae) pterygoid sinuses: phylogenetic and functional implications.

High-resolution X-ray computed tomographic scans were used to examine pterygoid sinus morphology within extant porpoise species and one delphinid (Tursiops truncatus), in order to consider: 1) intraspecific and interspecific variation among the studied species; 2) the most parsimonious sequence of character acquisition; and 3) the potential functional roles of the preorbital lobes of the sinuses in sound reflection. Scans revealed that the pterygoid/palatine regions are mediolaterally broader in the earliest diverging phocoenid (Neophocaena phocaenoides) and Tursiops truncatus than the dorsoventrally elongated sinuses observed in other species. Rostrocaudal lengths of the sphenoidal regions of the sinuses in all individuals studied are proportionally similar, indicating conservatism in this region across species. The neonate Phocoena phocoena has shorter preorbital lobes than adults, but they are still proportionally longer than Neophocaena phocaenoides and Phocoena spinipinnis. The preorbital lobes broaden mediolaterally to varying degrees across species; in particular, Phocoenoides dalli has the largest dorsal and lateral expansion of this region. Assuming the highest pulse frequency produced by porpoises is 150 kHz, all regions of the preorbital lobes are thick enough to reflect the wavelengths produced. In addition, the neonate preorbital lobes are not as elongated as they are in adults, and the dorsal third of this region may not reflect sound to the same extent. This study reinforces the importance of using nondestructive methods to quantify variation in endocranial anatomy and the value of CT data for recovering phylogenetically useful information, as well as functional roles sinuses play in concert with the soft tissue head anatomy for biosonar.

Hyperbaric computed tomographic measurement of lung compression in seals and dolphins.

Lung compression of vertebrates as they dive poses anatomical and physiological challenges. There has been little direct observation of this. A harbor and a gray seal, a common dolphin and a harbor porpoise were each imaged post mortem under pressure using a radiolucent, fiberglass, water-filled pressure vessel rated to a depth equivalent of 170 m. The vessel was scanned using computed tomography (CT), and supported by a rail and counterweighted carriage magnetically linked to the CT table movement. As pressure increased, total buoyancy of the animals decreased and lung tissue CT attenuation increased, consistent with compression of air within the lower respiratory tract. Three-dimensional reconstructions of the external surface of the porpoise chest showed a marked contraction of the chest wall. Estimation of the volumes of different body compartments in the head and chest showed static values for all compartments except the lung, which showed a pressure-related compression. The depth of estimated lung compression ranged from 58 m in the gray seal with lungs inflated to 50% total lung capacity (TLC) to 133 m in the harbor porpoise with lungs at 100% TLC. These observations provide evidence for the possible behavior of gas within the chest of a live, diving mammal. The estimated depths of full compression of the lungs exceeds previous indirect estimates of the depth at which gas exchange ceases, and concurs with pulmonary shunt measurements. If these results are representative for living animals, they might suggest a potential for decompression sickness in diving mammals.

Interspecific variation of ontogeny and skull shape among porpoises (Phocoenidae).

All extant members of Phocoenidae (porpoises) have been characterized as pedomorphic based on skeletal characters. To investigate the ontogenetic background for pedomorphosis and assess interspecific differences in ontogeny among phocoenids, samples of the six extant species were compared in terms of development of both epiphyseal and cranial suture fusion. Across all species, full maturity of the vertebral column was rare. Vertebral epiphyseal development did not progress so far in most Phocoena phocoena as in Phocoenoides dalli and Phocoena dioptrica. P. phocoena, Phocoena spinipinnis, Ph. dalli, and P. dioptrica, for which large series were available, were further compared in terms of ontogeny of cranial shape by three-dimensional geometric morphometrics. Ph. dalli and P. dioptrica generally showed further development of cranial sutures than the other species. Postnatal skull shape development was similar for all species studied; the majority of interspecific shape differences are present at parturition. Smaller species had a higher rate of shape development relative to growth in size than Ph. dalli and P. dioptrica, but they still showed less allometric development due to less postnatal growth. Interspecific shape differences indicate phylogenetic relationships similar to that proposed based on morphology or convergent evolution of the two pelagic species, Ph. dalli and P. dioptrica, under the scenarios suggested by recent molecular studies. A shape trend coinciding with habitat preference was detected; in species with pelagic preference the position and orientation of the foramen magnum aligned the skull with the vertebral column; the rostrum showed less ventral inclination, and the facial region was larger and more concave in lateral aspect.

B-mode ultrasonographic evaluation of the testis in relation to serum testosterone concentration in male Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis) during the breeding season.

The use of ultrasonography as a noninvasive tool for assessing the reproductive status of the male Yangtze finless porpoise (YFP; Neophocaena phocaenoides asiaeorientalis) was validated by correlating ultrasonographically determined testicular volume (TV) and testicular parenchyma pixel intensity (PI) with serum testosterone (T) concentration. The testes of 13 free-ranging male YFPs from the Tian-e-Zhou Reserve and three captive animals from the Baiji Dolphinarium (Wuhan, China) were examined ultrasonographically during April 2008. Testis volume was determined using Lambert's formula for an ellipsoid. Testicular parenchyma PI was evaluated by analyzing testicular ultrasonograms using pixel analysis software (Image J). Serum T concentrations were determined using a single-antibody radioimmunoassay. The TV, PI, and serum T concentration were low and similar in animals with body length <133 cm, highest in those with body length >or=142 cm, and highly variable in those with body length from 133 to 141 cm. Both TV and PI were closely correlated with serum T concentration (r=0.91 and r=0.85, respectively; P<0.01), indicating a consistent association between structural and functional development of the testis. In conclusion, we inferred that puberty onset in male YFPs occurred when TV was >150 cm(3) and PI was >60 during the breeding season and that testicular ultrasonography and pixel analysis was an efficient, noninvasive, real-time tool to evaluate testicular function of live male YFPs.

Ultrastructure of the spermatozoa of the Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis).

Semen sample was collected from two captive adult Yangtze finless porpoises (Neophocaena phocaenoides asiaeorientalis) during physical examination. One individual was aged about 9 years with body length 143 cm (total length) and body weight 46.1 kg in 2003. The age of the other was unknown and its body length was 147 cm and body weight was 43 kg in 2004. Ultrastructure of their spermatozoa was examined using scanning and transmission electron microscope. The sperm concentration was 4.17 x 10(9) spermatozoa per ml by the cytometer. The approximate dimensions of the spermatozoa were as follows: head length, 3.366 +/- 0.140 microm (mean +/- SE, n = 15); head width, 1.896 +/- 0.099 microm (n = 15); and neck length, 1.004 +/- 0.074 microm (n = 10). The tail included midpiece, principal piece and terminal piece. The length of the midpiece was 1.882 +/- 0.077 microm (n = 9). There is no apparent boundary between the principal piece and the terminal piece, so the length of the principal piece and the terminal piece was 44.612 +/- 3.485 microm (n = 5). Total length of the spermatozoa was 53.314 +/- 4.580 microm (n = 10). The acrosome covered approximately 45.8% of the anterior portion of the head.

Detecting recent speciation events: the case of the finless porpoise (genus Neophocaena).

Recent speciation events provide important insights into the understanding and conservation of Earth's biodiversity, representing recent adaptations to a changing environment and an important source of future evolutionary potential. However, the most frequently applied criterion for molecular-based speciation investigations, that of reciprocal monophyly of mitochondrial sequences, overlooks recent speciation events where insufficient time has passed for fixed molecular differences to develop between putative species. Two morphologically distinguishable forms of finless porpoise (genus Neophocaena) exist in sympatry in the strait of Taiwan, however the taxonomic relationship of these different forms is controversial. To test the hypothesis that the two forms represent different species, a study was conducted based on morphological characters and microsatellite and mitochondrial markers. The data suggest that the two forms are highly differentiated in terms of both morphology and genetic characteristics, despite being sympatric, and therefore represent different species as defined by the biological species concept. Moreover, the two forms appear to have been reproductively isolated since sharing a common ancestor prior to the last major glaciation event approximately 18 000 years ago. However, this represents an insufficient amount of time for reciprocal monophyly to have developed, and thus previous studies based on this criterion have overlooked this speciation event and resulted in incorrect taxonomic classification of these forms.

Comparative anatomy of the foramen ovale in the hearts of cetaceans.

The structure of the cardiac foramen ovale from 17 species representing six cetacean families, the Monodontidae, Phocoenidae, Delphinidae, Ziphiidae, Balaenidae and the Balaenopteridae, was studied using the scanning electron microscope. Eight white whale fetuses (Delphinapterus leucas) and a narwhal fetus (Monodon monoceros) represented the Monodontidae; one fetal and nine neonatal harbour porpoises (Phocoena phocoena) and a finless porpoise fetus (Neophocoena phocoenoides) represented the Phocoenidae; two white-beaked dolphin fetuses (Lagenorhynchus albirostris), four fetal and one neonatal Atlantic white-sided dolphins (Lagenorhynchus acutus), a Risso's dolphin fetus (Grampus griseus), two common bottle-nosed dolphin neonates (Tursiops truncatus), a female short-beaked common dolphin fetus (Delphinus delphis), four killer whale fetuses (Orcinus orca) and two long-finned pilot whale fetuses (Globicephala melas) represented the Delphinidae; two northern bottlenose whale fetuses (Hyperoodon ampullatus) represented the Ziphiidae; one bowhead whale fetus (Balaena mysticetus) represented the Balaenidae and five Common minke whale fetuses (Balaenoptera acutorostrata), one blue whale fetus (Balaenoptera musculus), nine fin whale fetuses (Balaenoptera physalus) and four humpback whale fetuses (Megaptera novaeangliae) represented the Balaenopteridae. The hearts of an additional two incompletely identified toothed and four baleen whale fetuses were also studied. In each species the fold of tissue derived from the cardiac septum primum and subtended by the foramen ovale had the appearance of a short tunnel or sleeve which was fenestrated at its distal end. In the toothed whales the tissue fold was tunnel-shaped with the interatrial septum as the floor whereas in baleen whales it was more sleeve-like. In toothed whales thin threads extended from the fold to insert into the interatrial septum whereas a network of threads covered the distal end of the sleeve in the baleen whales. Similar structures were present in the corresponding cardiac tissues of neonatal Hippopotamidae.

Structure of the cerebral cortex of the humpback whale, Megaptera novaeangliae (Cetacea, Mysticeti, Balaenopteridae).

Cetaceans diverged from terrestrial mammals between 50 and 60 million years ago and acquired, during their adaptation to a fully aquatic milieu, many derived features, including echolocation (in odontocetes), remarkable auditory and communicative abilities, as well as a complex social organization. Whereas brain structure has been documented in detail in some odontocetes, few reports exist on its organization in mysticetes. We studied the cerebral cortex of the humpback whale (Megaptera novaeangliae) in comparison to another balaenopterid, the fin whale, and representative odontocetes. We observed several differences between Megaptera and odontocetes, such as a highly clustered organization of layer II over the occipital and inferotemporal neocortex, whereas such pattern is restricted to the ventral insula in odontocetes. A striking observation in Megaptera was the presence in layer V of the anterior cingulate, anterior insular, and frontopolar cortices of large spindle cells, similar in morphology and distribution to those described in hominids, suggesting a case of parallel evolution. They were also observed in the fin whale and the largest odontocetes, but not in species with smaller brains or body size. The hippocampal formation, unremarkable in odontocetes, is further diminutive in Megaptera, contrasting with terrestrial mammals. As in odontocetes, clear cytoarchitectural patterns exist in the neocortex of Megaptera, making it possible to define many cortical domains. These observations demonstrate that Megaptera differs from Odontoceti in certain aspects of cortical cytoarchitecture and may provide a neuromorphologic basis for functional and behavioral differences between the suborders as well as a reflection of their divergent evolution.

Sexually dimorphic proportions of the harbour porpoise (Phocoena phocoena) skeleton.

Sexual differences in growth, allometric growth patterns and skeletal proportions were investigated by linear measurements of skeletal parts on 225 harbour porpoises (Phocoena phocoena) from the inner Danish and adjacent waters. Females show larger asymptotic sizes and extended period of growth compared with males. Measurements of the skull and flipper bones show negative allometry, whereas those of the bones of the body generally show positive allometry. There are no statistically significant intersexual differences in allometry except for the pelvic bones, where the males show stronger positive allometry. Throughout the range of individual sizes, females have significantly larger skulls and shorter vertebral columns than males for similarly sized individuals. In fully grown specimens, the condylobasal length of females makes up a smaller proportion of total length, and the vertebrae make up a larger proportion as compared with males. As these characters show negative and positive allometry, respectively, it is suggested that males finish their development at an earlier stage than females, retaining more paedomorphic proportions of the skeleton. Paedomorphosis in fully grown males relative to females is also found in the vertebral epiphyses that mature later in males than females, although the males finish growth at a younger age.

Comparative anatomical study on the relationships between the vestigial pelvic bones and the surrounding structures of finless porpoises (Neophocaena phocaenoides).

Morphology of the modern cetaceans represents the results of adaptation of the ancestral terrestrial mammals to aquatic life through their evolutional processes. Some of the primitive fossil cetaceans are known to have both fore and hind limbs, whereas the pelvic bones of modern cetaceans are, in general, a pair of slender rod-like structures within the abdominal wall muscles just anterior to the anus with no articulations to the axial skeleton in both sexes. It is interesting and important to consider the causes and processes of how the hind limbs were lost and how the pelvis was reduced during the process of adaptation. In the present study, we tried to evaluate the topography and function of rudimentary pelvic bones of the finless porpoise (Neophocaena phocaenoides), one of the members of the odontocete cetaceans, with special references to the structures around the pelvic bones. Some soft tissues such as M. ischiocavernosus relating to the pelvic bone are transformed following the drastic reduction of the pelvis. This transformation tells us that the cetaceans adapted to the aquatic life during evolutional processes chose the tail flukes driven by the powerful trunk muscles for locomotion, instead of modifying the hind limbs into hind flippers as seen in pinnipeds. On the other hand, it is evident that a function of the pelvic bones of the male finless porpoise was supporting the penis as those of terrestrial mammals. It is noteworthy that the morphological features of the ancestral terrestrial mammals can be traced when they are carefully compared with those of the finless porpoise.

Structural fiber reinforcement of keel blubber in harbor porpoise (Phocoena phocoena).

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.

Patterns of ossification in the manus of the harbor porpoise (Phocoena phocoena): hyperphalangy and delta-shaped bones.

With the transition from terrestrial to aquatic habitats, cetacean forelimbs have undergone significant modifications in bone morphology and soft tissue distribution. Some, but not all, of these modifications are also demonstrated in other lineages of extant and extinct secondarily aquatic tetrapods. This study examines the ontogenetic pattern of ossification of the manus of the harbor porpoise (Phocoena phocoena), using plain film radiography. Two modifications examined are hyperphalangy (number of phalanges per digit increased beyond the typical mammalian number) and the morphology of delta-shaped bones. Hyperphalangy in Phocoena phocoena is apparent in digits 2 and 3. Phalangeal counts in all digits are variable (sometimes between the right and left flippers of the same individual) and are not necessarily correlated with age. Phalangeal ossification and epiphyseal fusion proceeds along the proximo-distal axis within each digit. In addition, digits 2 and 3 are at a more advanced stage of ossification than more abaxial digits. Delta-shaped bones appear to be a normal stage in the ossification of phalanges in all digits except the third, and may persist in the adult in certain digits. In humans, this morphology is a developmental anomaly usually associated with other malformations, such as polydactyly or syndactyly. Delta-shaped bones in the cetacean manus display a consistent orientation and the process by which they are formed may be similar to that in extinct marine reptiles.

Wing design and morphology of the harbor porpoise dorsal fin.

The correlation between skin structure and hydrodynamic design of the dorsal fin of the harbor porpoise (Phocoena phocoena) was examined. For the study of fin morphology and geometry, a scheme of sampling representing a two-parameter mesh on the fin surface was used. At each data point the thickness of the epidermis, papillary and subpapillary layers of the dermis, the ligamentous layer of the fin, as well as the angle formed by the direction of dermal ridges and the fin root chord were measured. On the basis of fin cross-sections the three-dimensional surface models of the fin in a 1 : 1 scale were created with a CAD program. The shape of the model was evaluated by the wing and hydrofoil parameters (angle of leading edge sweep, leading edge radius, maximum thickness of the fin cross-section, and position of maximum thickness from the leading edge). Hydrodynamic performance of the fin cross-sections was studied with a CFD program. Regional variability of the parameters of morphology was compared with spanwise variability of the parameters of cross-sectional geometry. It was found that skin structure parameters correlate with the hydrodynamically relevant parameters of the fin and fin cross-sections. Regularities of skin structure of the harbor porpoise dorsal fin are considered indirect evidence of the adaptation of porpoise skin to the fin flow.

Neuroanatomy of the harbor porpoise (Phocoena phocoena) from magnetic resonance images.

Cetacean (dolphin, whale, and porpoise) brains are among the least-studied mammalian brains because of the formidability of collecting and histologically preparing such relatively rare and large specimens. Among cetaceans, there exist relatively few studies of the brain of the harbor porpoise (Phocoena phocoena). Magnetic resonance imaging (MRI) offers a means of observing the internal structure of the brain when traditional histological procedures are not practical. Therefore, MRI has become a critical tool in the study of the brain of cetaceans and other large species. This article represents the first MRI-based anatomically labeled three-dimensional description of the harbor porpoise brain. Coronal plane sections of the brain of a young harbor porpoise were originally acquired and used to produce virtual digital scans in the other two orthogonal spatial planes. A sequential set of images in all three planes has been anatomically labeled and displays the proportions and positions of major neuroanatomical features. These images allow for the visualizing of the distinctive features of the harbor porpoise brain from various orientations by preserving the gross morphological structure of the specimen.

Edward Tyson's 1680 account of the 'porpess' brain and its place in the history of comparative neurology.

Edward Tyson (1650-1708), a prominent London physician and an early fellow of the Royal Society is best known for his several anatomical contributions in the creation of primatology, including the preputial and coronal mucilaginous glands (Tyson's glands), later described by Alexis Littré. He also published the first comprehensive account of a single animal (the 'porpess') and placed it in the context of a systematic and experimental methodology. This rare monograph accounts for the contention that Tyson was the founder of comparative anatomy in England, by using this 'fish' to better understand the human condition. His description of the highly convoluted cetacean brain as well as his recognition of the many homologies with "land-quadrupeds", rather than the fishes it resembled, constitutes a major landmark contribution to the history of biology. The prevailing theological thrust of training in 'physick' ultimately led Tyson to evade the conundrum of how the human brain differs from that of animals, by attributing the intellectual uniqueness of man to endowments derived from God rather than the physical substance of the brain.

Changes in blubber distribution and morphology associated with starvation in the harbor porpoise (Phocoena phocoena): evidence for regional differences in blubber structure and function.

To examine patterns of blubber loss accompanying a decline in body condition, blubber thickness of juvenile harbor porpoises in normal/robust body condition (n=69) was compared with that of starved conspecifics (n=31). Blubber thickness in the thorax of starved porpoises (9-11 mm) was only 50%-60% of that of normal animals (18-20 mm); however, very little tailstock blubber was lost during starvation. Adipocytes in thorax and tailstock blubber were measured in both groups (n=5) to determine whether thickness changes were homogeneous throughout blubber depth. In the thorax of normal porpoises, adipocytes near the epidermis (outer blubber) were smaller (0.11 nL) than inner blubber adipocytes (0.17 nL). Conversely, the size of tailstock adipocytes was uniform. Starved animals had fewer, smaller adipocytes in the inner thorax blubber, suggesting a possible combination of adipocyte shrinkage and loss. Lipids were withdrawn only from the inner layer of thorax blubber during starvation, supporting a hypothesis of regional specialization of function in blubber. Blubber of the thorax serves as the site of lipid deposition and mobilization, while the tailstock is metabolically inert and likely important in locomotion and streamlining. Therefore, some proportion of the blubber of small odontocetes must be considered structural/mechanical rather than an energy reserve.