SOME ANIMAL ACOUSTIC PROPERTIES. 2. OF ECHOES AND THEIR USES.
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A complex lymphoepithelial gland is a constant feature in the larynx of Atlantic bottlenose dolphins, Tursiops truncatus, based on study of 56 animals. Larynges were removed from fresh, non-decomposed beach-stranded animals for gross examination and histological sampling. A large lymphoepithelial gland occurs in the rostro-ventral mucosa of the larynx, overlying the cricoid cartilage. It presents as a well-defined, elevated, and heavily trabeculated area. Histological examination reveals a pseudostratified columnar epithelium which branches into the underlying submucosa. The epithelial-lined folds and crypts thus formed are surrounded by aggregations of lymphocytes, which infiltrate this epithelium. Mucous glands are often associated with these lymphoid aggregations. The histological appearance of the laryngeal gland is remarkably similar to the palatine, or dorsal oropharyngeal tonsils, of T. truncatus. It may be analogous to the nasopharyngeal adenoid of terrestrial animals. Age-related involution of the laryngeal gland is not as obvious with increasing animal age (or length) as it is in other mucosa-associated lymphoid tissue. The distribution of this gland among cetaceans is not yet known. We have observed it in individuals of every species we have studied, including Lagenodelphis hosei, Stenella coeruleoalba, Stenella attenuata, Globicephala macrorhynchus, Steno bredanensis, Physeter catodon, Pseudorca crassidens, Mesoplodon europaeus, and Kogia breviceps.
The site and physiologic mechanism(s) responsible for the generation of odontocete biosonar signals have eluded investigators for decades. To address these issues we subjected postmortem toothed whale heads to interrogation using medical imaging techniques. Most of the 40 specimens (from 19 species) were examined using x-ray computed tomography (CT) and/or magnetic resonance imaging (MR). Interpretation of scan images was aided by subsequent dissection of the specimens or, in one case, by cryosectioning. In all specimens we described a similar tissue complex and identified it as the hypothetical biosonar signal generator. This complex includes a small pair of fatty bursae embedded in a pair of connective tissue lips, a cartilaginous blade, a stout ligament, and an array of soft tissue air sacs. Comparing and contrasting the morphologic patterns of nasal structures across species representing every extant odontocete superfamily reveals probable homologous relationships, which suggests that all toothed whales may be making their biosonar signals by a similar mechanism.
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The heart from a 4-m-long minke whale (Balaenoptera acutorostrata) was studied to determine the details of its anatomy and to consider cardiac adaptations to diving. The volume fraction (Vvc) of capillaries in the wall of the left ventricle was determined at different levels from base to apex and at different depths from epi- to endocardium using a light microscopic stereologic technique. Typical of cetaceans, this minke whale heart was distinctly flattened dorsoventrally. A moderator band, characteristic of ungulate hearts, spanned the right ventricle. The right and left atrioventricular valves were tricuspid and bicuspid, respectively. The right coronary artery supplied the dorsal and right lateral myocardium. The left coronary artery supplied the ventral and left lateral myocardium. An anastomosis between the dorsal and ventral interventricular arteries occurred in the dorsal interventricular groove. Stereologically, a decreasing transmural gradient in Vvc was identified between the epicardium and the subepicardium at 15 cm from the apex. Our results, however, did not reveal any significant deviations in the pattern of capillary distribution in the wall of the left ventricle between this baleen whale and terrestrial mammals. Measurements of the heart, great vessels, coronary vasculature, and ventricular walls are also given, and they suggest a physiologic and adaptive right ventricular hypertrophy. Based on these and other observations, we propose that the relatively great thickness of the right ventricle and the distinctive shape of the cetacean heart are adaptations to the hemodynamic changes and collapse of the thorax associated with apneic diving.
In terrestrial mammals, stratum corneum lipids derive from two sources: deposition of lamellar body lipids in stratum corneum interstices and excretion of sebaceous lipids onto the skin surface, resulting in a two-compartment ("bricks and mortar") system of lipid-depleted cells surrounded by lipid-enriched intercellular spaces. In contrast, intracellular lipid droplets, normally not present in the epidermis of terrestrial mammals, are prominent in avian and marine mammal epidermis (cetaceans, manatees). We compared the transepidermal water loss, ultrastructure, and lipid biochemistry of the viable epidermis and stratum corneum of pigeon apterium, fledgling (featherless) zebra finches, painted storks, cetaceans, and manatees to those of humans and mice. Marine mammals possess an even more extensive lamellar-body secretory system than do terrestrial mammals; and lamellar-body contents, as in terrestrials, are secreted into the stratum corneum interstices. In cetaceans, however, glycolipids, but not ceramides, persist into the stratum corneum; whereas in manatees, glycolipids are replaced by ceramides, as in terrestrial mammals. Acylglucosylceramides, thought to be critical for lamellar-body deposition and barrier function in terrestrial mammals, are present in manatees but virtually absent in cetaceans, a finding that indicates that they are not obligate constituents of lamellar-body-derived membrane structures. Moreover, cetaceans do not elaborate the very long-chain, saturated N-acyl fatty acids that abound in terrestrial mammalian acylglucosylceramides. Furthermore, cold-water marine mammals generate large, intracellular neutral lipid droplets not found in terrestrial and warm-water marine mammals; these lipid droplets persist into the stratum corneum, suggesting thermogenesis, flotation, and/or cryoprotectant functions. Avians generate distinctive multigranular bodies that may be secreted into the intercellular spaces under xerotic conditions, as in zebra fledglings; ordinarily, however, the internal lamellae and limiting membranes deteriorate, generating intracellular neutral lipid droplets. The sphingolipid composition of avian stratum corneum is intermediate between terrestrials and cetaceans (approximately equal to 50% glycolipids), with triglycerides present in abundance. In the midstratum corneum of avians, neutral lipid droplets are released into the interstices, forming a large extracellular, lipid-enriched compartment, surrounding wafer-thin corneocytes, with a paucity of both lipid and keratin ("plates-and-mortar" rather than the "bricks-and-mortar" of mammals).(ABSTRACT TRUNCATED AT 400 WORDS)
Gastric and cranial duodenal structure of the bowhead whale (Balaena mysticetus) was examined grossly and microscopically. The stomach was arranged in a series of four compartments. The first chamber, or forestomach, was a large nonglandular sac lined by a keratinized stratified squamous epithelium. It was followed by the fundic chamber, a large, somewhat globular and entirely glandular compartment. At the entrance of the fundic chamber, a narrow cardiac gland region could be defined. The remaining mucosa of the chamber contained the proper gastric glands. A narrow, tubular connecting channel, the third distinct gastric division, was lined by mucous glands and joined the fundic chamber with the final stomach compartment, or pyloric chamber. This fourth chamber was also tubular and lined by mucous glands but was of a diameter considerably larger than the connecting channel. The stomach terminated at the pyloric sphincter which consisted of a well-developed band of circular smooth-muscle bundles effecting a division between the pyloric chamber and small intestine. The small intestine began with the duodenal ampulla, a dilated sac considerably smaller than the fundic chamber of the stomach. The mucosa of this sac contained mucous glands throughout. The ampulla led without a separating sphincter into the duodenum proper which continued the intestine in a much more narrow tubular fashion. The mucosal lining of the duodenum was composed of villi and intestinal crypts. Although their occurrence varied among whales, enteroendocrine cells were identified within the mucous glands of the cardiac region, connecting channel, pyloric chamber, and cranial duodenum. The hepatopancreatic duct entered the wall of the duodenum shortly after the termination of the duodenal ampulla and continued intramurally along the intestine before finally joining the duodenal lumen.
A late Miocene/early Pliocene mammalian fossil from the site of Sahabi, Libya, was identified in previous publications as a clavicle belonging to the superfamily Hominoidea. After studying both a cast of the specimen and previously published descriptions and photographs we have concluded that the fossil is not a clavicle, but a rib fragment. In addition, our study demonstrates beyond reasonable doubt that this rib belongs not to a primate, but to a marine mammal. Following comparisons with 19 species in 14 genera of marine mammals, as well as six hominoid and three cercopithecoid species, both modern and fossil, we conclude that the specimen is best attributed to the Odontoceti, the cetacean suborder containing dolphins and porpoises. The rib indicates an animal in the size range of Recent Tursiops or Lagenorhynchus.
The ratios of 15N to 14N and 13C to 12C tend to be higher in marine than in terrestrial organisms. The concentrations of these isotopes in human bone collagen consequently can be used to make inferences about the contribution of marine and terrestrial resources to prehistoric diets. The utility of studying 15N/14N and 13C/12C ratios in conjunction with each other is illustrated by our analysis of 40 human burials from archaeological sites in the Santa Barbara Channel area of southern California. The mean delta 13C and delta 15N values (in per mil) of collagen from these skeletons decrease progressively from the Channel Islands (delta 13C = -14.0, delta 15N = +16.3) to the mainland coast (delta 13C = -14.5, delta 15N = +14.9) to the interior (delta 13C = -17.2, delta 15N = +10.9). These data suggest that Indians living on the Channel Islands during the late prehistoric period were heavily dependent on marine resources. The inhabitants of the mainland interior, in contrast, had a diet composed largely of terrestrial foods. From their isotope ratios, it appears that the Indians who lived on the mainland coast consumed a mixed diet containing substantial quantities of both marine and terrestrial resources. Differences in 15N/14N and 13C/12C ratios of individuals from mainland sites dating from the early and late prehistoric periods show that the marine component of the diet increased substantially through time. These isotopic data are consistent with pathological, faunal, and artifactual evidence of increased marine resource exploitation during the late prehistoric period.
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Components of the respiratory system from seven bowhead whales have been examined. The paired and laterally curved external nares are passively closed by a valve-like mass located in the rostral, lateral, and ventral walls of the nasal vestibules. Nasal septal cartilages are paired smooth plates rostrally changing to accordion-like folds caudally. The epiglottic and arytenoidal protuberances of the larynx are typically cetacean, but blunt. The cricoid cartilage is not a complete ring, but an elongated, inverted, trough-shaped structure. The thyroid cartilage is trough-shaped with elongated cranial cornua curving dorsocaudally from each thyroid lamina. A conical mass of skeletal muscle serves as the floor of the short trachea and also surrounds the termination of the laryngeal sac. The trachea is dorsoventrally compressed, lacks a tracheal bronchus, and its width equals its length. The principal bronchi give rise to lobar bronchi at obtuse angles. Large segmental bronchi branch extensively from lobar bronchi near the mediastinal lung surface. The lungs are rectangular and of nearly uniform thickness throughout, without external or internal lobulation.
The macroanatomy of renicules and surrounding tissues from the kidneys of five Eskimo-harvested bowhead whales, Balaena mysticetus , was examined. These renicules are similar in overall structure to those of other cetaceans and intermediate in size. There are several important differences including the presence of arcuate vessels within the sporta perimedullaris , the extension of connective tissue from the sporta deep into the peripheral cortex, and the presence of very large, thin-walled veins that occupy the interrenicular spaces. Arterial and venous plexuses outside the substance of the sporta reported in other cetaceans were not observed in the bowhead .
In the bowhead whale, Balaena mysticetus, the mucosa of the major airways from the blowholes through the rostral portion of the larynx is lined with parakeratotic, pigmented, stratified squamous epithelium. Scattered enlarged connective tissue papillae of the lamina propria of the nasal vestibules and the palatopharyngeal sphincter contain encapsulated nerve endings. Abundant papillae in the mucosa covering the epiglottic and arytenoid cartilages contain similar nerve endings. The remainder of the laryngeal cavity and laryngeal sac is lined by a variably pigmented, stratified squamous epithelium, which is not keratinized. At the laryngotracheal junction the lining changes to ciliated pseudostratified columnar epithelium which continues through the trachea and principal bronchi. Scanning electron microscopy (SEM) indicates that this epithelium is typically mammalian, with approximately half of the surface cells bearing cilia and slender microvilli. The remaining cells are mucus producing and have thicker microvilli. The valvular mass regulating the external nares consists of irregular, dense white fibrous connective tissue with numerous adipose cells and is penetrated by skeletal muscle cords ranging from 2-4 mm in diameter. The septal mass between the blowholes is composed of irregular, dense white fibrous connective tissue containing large tendinous bundles, clusters of adipose cells, and several large arteries and thick-walled veins. The lamina propria of the nasal vestibules is irregular, dense white fibrous connective tissue. That of the larynx is not as dense and contains proportionately more elastic fibers. The laryngeal sac does not contain elastic laminae, but does have a tunica muscularis of skeletal muscle bundles. Within the trachea and principal bronchi, the lamina propria possesses laminae of longitudinally oriented elastic fibers and simple, branched tubuloalveolar mucous glands. The nasal, laryngeal, tracheal, and bronchial cartilages are hyaline with vascular channels.
Skin samples from most body regions of the bowhead whale were examined. The epidermis is 2.7 to 50 times thicker than that reported in other cetaceans with both regional and individual variations in thickness. The thinnest areas examined (1 mm) occur on the eyelid margins and the thickest (25 mm) occur on the lower jaw. A distinctive parakeratotic stratum corneum with a thick underlying stratum spinosum (without a stratum granulosum) extends over the entire body surface. From a few dozen to several hundred epidermal lesions are present on all whales studied. A typical stratum basale of germinative keratinocytes (with melanocytes in pigmented areas) rests upon a well-defined basal lamina. Epidermal rod arrays arise from the basal keratinocytes which cover highly elongated dermal papillae and extend to the epidermal surface through the distal stratum spinosum and the stratum corneum. At least four diatom genera occur on and in the stratum corneum and lesion areas of different whales. The superficial dermis consists of a papillary layer with long (up to 13 mm) dermal papillae interdigitating with the epidermis from a basal area that is 2-4 mm in thickness. The number of dermal papillae per mm2 varies inversely with the thickness of the epidermis. Large diameter, sensory papillae packed with tortuous, highly elongated, encapsulated nerve end organs also interdigitate with the thin epidermal areas of the ventral surface of the rostrum, the upper and lower lip margins, and the upper and lower eyelid margins. Scattered, single, stiff hairs emerge from the skin only in specific, pigmented regions of the head.