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[The adaptation to water of whale skin. Histologic and histochemical studies of the dolphin, Delphinus delphis, and the porpoise, Phocaena phocaena].

In the whale the superficial layer of the epidermis (stratum paracorneum) is incompletely cornified and is characterized by high contents of phospholipids, neutral fats, and glycoproteins. This finding, and especially the structure of the dermis (hypodermis) in relation to its function, are discussed as to the role they play in the adaptation of whales to the aquatic environment.

Adaptation, Biological↗

Source-to-sensation level ratio of transmitted biosonar pulses in an echolocating false killer whale.

Transmitted biosonar pulses, and the brain auditory evoked potentials (AEPs) associated with those pulses, were synchronously recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP amplitude was investigated as a function of the transmitted biosonar pulse source level. For that, a few thousand of the individual AEP records were sorted according to the spontaneously varied amplitude of synchronously recorded biosonar pulses. In each of the sorting bins (in 5-dB steps) AEP records were averaged to extract AEP from noise; AEP amplitude was plotted as a function of the biosonar pulse source level. For comparison, AEPs were recorded to external (in free field) sound pulses of a waveform and spectrum similar to those of the biosonar pulses; amplitude of these AEPs was plotted as a function of sound pressure level. A comparison of these two functions has shown that, depending on the presence or absence of a target, the sensitivity of the whale's hearing to its own transmitted biosonar pulses was 30 to 45 dB lower than might be expected in a free acoustic field.

Acoustics↗

The neural network classification of false killer whale (Pseudorca crassidens) vocalizations.

This study reports the use of unsupervised, self-organizing neural network to categorize the repertoire of false killer whale vocalizations. Self-organizing networks are capable of detecting patterns in their input and partitioning those patterns into categories without requiring that the number or types of categories be predefined. The inputs for the neural networks were two-dimensional characterization of false killer whale vocalization, where each vocalization was characterized by a sequence of short-time measurements of duty cycle and peak frequency. The first neural network used competitive learning, where units in a competitive layer distributed themselves to recognize frequently presented input vectors. This network resulted in classes representing typical patterns in the vocalizations. The second network was a Kohonen feature map which organized the outputs topologically, providing a graphical organization of pattern relationships. The networks performed well as measured by (1) the average correlation between the input vectors and the weight vectors for each category, and (2) the ability of the networks to classify novel vocalizations. The techniques used in this study could easily be applied to other species and facilitate the development of objective, comprehensive repertoire models.

Animals↗

Characterizing the graded structure of false killer whale (Pseudorca crassidens) vocalizations.

The vocalizations from two, captive false killer whales (Pseudorca crassidens) were analyzed. The structure of the vocalizations was best modeled as lying along a continuum with trains of discrete, exponentially damped sinusoidal pulses at one end and continuous sinusoidal signals at the other end. Pulse trains were graded as a function of the interval between pulses where the minimum interval between pulses could be zero milliseconds. The transition from a pulse train with no inter-pulse interval to a whistle could be modeled by gradations in the degree of damping. There were many examples of vocalizations that were gradually modulated from pulse trains to whistles. There were also vocalizations that showed rapid shifts in signal type--for example, switching immediately from a whistle to a pulse train. These data have implications when considering both the possible function(s) of the vocalizations and the potential sound production mechanism(s). A short-time duty cycle measure was developed to characterize the graded structure of the vocalizations. A random sample of 500 vocalizations was characterized by combining the duty cycle measure with peak frequency measurements. The analysis method proved to be an effective metric for describing the graded structure of false killer whale vocalizations.

Animals↗

Hemolytic plaque inhibition by synthetic antigenic peptides of sperm whale myoglobin.

We have investigated the systemic antibody response to sperm whale myoglobin (SWMb) antigenic sites in three strains of inbred mice using an inhibition of plaque assay. Sperm whale myoglobin was attached to sheep red blood cells (SRBC) via rabbit anti-SRBC Fab' fragments. Inhibition of lysis was obtained with synthetic peptides representing the purported five antigenic sites but not with a peptide whose sequence was unrelated to SWMb and synthetic peptides of SWMb from outside the antigenic sites gave minimal or no inhibition. The results of our studies show that the pattern of response to the five antigenic sites differs in each strain, but that almost total inhibition is obtained in all strains with these five sites. The antigenic dominance of these sites supports the concept of discrete antigenic sites on soluble proteins. They also suggest a reason for contrary reports in the literature based on hybridoma technology.

Animals↗

[Supply of medicines for the Dutch whaling and salt trade in de seventeenth century].

During the seventeenth century the small Dutch nation with a population of no more than 1.5 million people played a prominent role in the whale oil and salt trade. Two reports discovered in the Archives in The Hague and Delft are dealing with the provision of medical and pharmaceutical services during their extensive maritime commerce. Surgeon chests destinated for whaling explicitly contained medicines to treat scurvy, respiratory illnessses and musculoskeletal diseases. The workers exploiting the saltpans in Arguin, near the northwestern coast of Afrika, were in need for drugs to cure gastrointestinal complaints and inflammatory conditions of the mouth.

Commerce↗

The whale forgetting factor in recursive AR spectral analysis of heart rate variability signals.

Spectral parameters extracted from the heart rate variability signal are obtained on a beat-to-beat basis by means of autoregressive recursive identification. In this paper a whale forgetting window is introduced, instead of the classical exponential one, in order to reduce the noise influence on the estimated parameters. After proper simulation it was found that the whale forgetting window markedly reduces the noise in the identification, but maintains a good response to abrupt changes in the signal. The algorithm was thus applied to the analysis of the HRV data recorded during different transient situations in physiological and pathological conditions. The spectral parameters were obtained on a beat-to-beat basis and their trends were smoother and more accurate with respect to the traditional exponential window also in presence of noise or artifacts in the time series (sudden and short time changes, ectopic beats, etc.), without losing the signal variations of physiological interest.

Adult↗

Anatomy and three-dimensional reconstructions of the brain of the white whale (Delphinapterus leucas) from magnetic resonance images.

Magnetic resonance imaging offers a means of observing the internal structure of the brain where traditional procedures of embedding, sectioning, staining, mounting, and microscopic examination of thousands of sections are not practical. Furthermore, internal structures can be analyzed in their precise quantitative spatial interrelationships, which is difficult to accomplish after the spatial distortions often accompanying histological processing. For these reasons, magnetic resonance imaging makes specimens that were traditionally difficult to analyze, more accessible. In the present study, images of the brain of a white whale (Beluga) Delphinapterus leucas were scanned in the coronal plane at 119 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 such structures as the corpus callosum, internal capsule, cerebral peduncles, cerebral ventricles, certain thalamic nuclear groups, caudate nucleus, ventral striatum, pontine nuclei, cerebellar cortex and white matter, and all cerebral cortical sulci and gyri.

Animals↗

Gross anatomy of the respiratory system of the bowhead whale, Balaena mysticetus.

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.

Animals↗

Macroanatomy of the renicule of the bowhead whale (Balaena mysticetus).

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 .

Animals↗

Microanatomy of the major airway mucosa of the bowhead whale, Balaena mysticetus.

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.

Animals↗

Position of the larynx in odontoceti (toothed whales).

This study examines the positional relationships of the odontocete (toothed whale) larynx to further an understanding of their breathing, swallowing, and vocalizing abilities. Seventeen specimens representing nine cetacean genera (Delphinus, Stenella, Tursiops, Grampus, Delphinapterus, Globicephala, Kogia, Mesoplodon, and Phocoena) were studied post mortem. Nine specimens were sectioned in the midsagittal plane and the position of the larynx relative to vertebral levels, skull base, and palatal structures was recorded. In eight specimens that could not be bisected for reasons of large size or condition of preservation, the larynx was removed by a ventral approach for further dissection. The results show that the upper respiratory tract of the odontocetes has evolved away from a basic mammalian pattern. Laryngeal position among terrestrial mammals usually corresponds to the level of cervical vertebrae 1-3. The odontocete larynx, however, lies rostral to the level of the atlas and extends to the presphenoidal synchondrosis. Its extension above the level of the foramen magnum is due to three factors: 1) The larynx is elongated into a tubular extension that projects beyond the soft palate into the nasopharynx; 2) the neck region is shortened owing to the highly compressed cervical vertebrae; and 3) the skull base is oriented in the same direction as the cervical vertebrae because of the horizontal and fusiform alignment of the head and thorax. Whereas the larynx of most terrestrial mammals is separable from the nasopharynx, that of the odontocetes studied may be permanently intranarial, held in place by the palatopharyngeal sphincter. Laryngeal position may affect their vocal abilities, allowing odontocetes to simultaneously swallow and echolocate.

Animals↗

Existence of vocal folds in the larynx of odontoceti (toothed whales).

Odontocetes (toothed whales) vocalize for communication and echolocation. The mechanisms of sound production, however, remain unclear. Their larynx has long been thought to lack vocal folds and, thus, was considered incapable of generating sounds. This study investigates internal anatomy of the odontocete larynx to: 1) describe the morphology of any folds found, 2) determine any structural homologies between these folds and the vocal folds of terrestrial mammals, and 3) assess their possible function in sound production. Larynges of 24 odontocetes representing ten genera (Delphinus, Stenella, Lagenorhynchus, Tursiops, Grampus, Delphinapterus, Globicephala, Kogia, Mesoplodon, and Phocoena) were studied post mortem. Nine specimens were cut midsagittally, and the remainder were dorsally opened to reveal internal anatomy. Results show that, contrary to established belief, vocal folds are consistently present. They are not isolated bands or "cords," but appear continuous with the internal laryngeal membrane. The attachments of these folds are the same as in terrestrial mammals, thus indicating homology with true mammalian vocal folds. These folds extend from the midline of the thyroid cartilage to the base of the arytenoid cartilages, sometimes to a discrete process. The vocal folds are elongated and oriented in a vertical plane, parallel to airflow direction. Vocal fold morphology varies, appearing as true bifurcated structures, a trifurcated fold, or a single midline fold. Laryngeal ventricles and vestibular folds are also consistently found lateral to the vocal folds. The vocal folds may divide the airstream within the larynx into three separate air currents. Fold vibrations may produce initial laryngeal sound used in echolocation or communication.

Animals↗

Microanatomy of the lung of the bowhead whale Balaena mysticetus.

The lungs from six bowhead whales harvested by Alaskan Eskimos have been examined with light and electron microscopes. Airways ranging from 1 to 40 mm in luminal diameter are lined by a pseudostratified ciliated epithelium containing numerous mucus-secreting cells. The underlying lamina propria-tela submucosa of these airways contains tubuloalveolar glands, plasma cells, and lymphatic accumulations in addition to both elastic and collagenous fibrillar elements. Cartilage extends to the level of the respiratory airways, but smooth muscle is absent from airways larger than 3 mm, and tubuloalveolar glands are absent from airways smaller than 3 mm. Respiratory airways are lined by pseudostratified, simple cuboidal, and simple squamous epithelia. Alveolar ducts are lined by simple squamous epithelium exclusively. A connective tissue core composed mostly of elastic fibers supports the walls of the alveolar ducts. Neither smooth muscle nor cartilage has been observed in these structures. Alveoli contain the typeical cetacean double capillary bed separated by a thick septum composed mainly of collagenous connective tissue. Alveoli are lined by a simple squamous epithelium similar to that encountered in alveolar ducts and respiratory airways. This epithelium is composed of type I and II pneumocytes closely appressed to an underlying capillary network. The type II pneumocytes contain typical lamellar bodies and tubular myelin can be seen in the air spaces. The lung is surrounded by a thick (X = 2.5 mm) visceral pleura rich in blood vessels and elastic fibers.

Animals↗

Anatomy of the hyoid apparatus in Odontoceti (toothed whales): specializations of their skeleton and musculature compared with those of terrestrial mammals.

BACKGROUND: The hyoid apparatus of odontocetes (toothed whales) serves as a major attachment point for many of the muscles and ligaments subserving breathing, swallowing, and sound production. METHODS: This study examines the hyoid apparatus in 48 specimens of ten odontocete genera (Phocoena, Lagenorhynchus, Stenella, Delphinus, Tursiops, Grampus, Globicephala, Mesoplodon, Physeter, and Kogia) collected post mortem from beach strandings. RESULTS: The odontocete hyoid apparatus, as that of their closest terrestrial relatives, the artiodactyls, is divisible into a basal portion (basihyal, paired thyrohyals) and a suspensory portion (paired ceratohyals, epihyals, stylohyals, and tympanohyals) connecting the basal portion to the skull base. Unlike other terrestrial mammals, the basal portion lies inferior to the laryngeal aditus, is flattened dorso-ventrally, and is relatively large, thus providing a broad surface area for muscle attachments. The suspensory elements are not as flattened and are joined by synovial joints (except for epihyal-stylohyal fusion). Muscular specializations include enlargement of those which retract the hyoid apparatus (e.g., sternohyoid) or control the tongue (e.g., styloglossus, hyoglossus). These muscles may be particularly important in a specialized prey capture behavior called suction feeding. In addition, the hyoid apparatus has a tilted placement, which allows asymmetrical enlargement of the piriform sinuses. Asymmetry is also seen in the muscular attachments between the larynx and the hyoid apparatus. The most pronounced differences from the basic pattern are observed in two families: Physeteridae and Ziphiidae. CONCLUSIONS: The derived position and shape of the odontocete hyoid apparatus may have evolved to subserve several specialized upper respiratory/digestive tract functions, such as simultaneous feeding (suction and swallowing) and sound production.

Anatomy, Comparative↗

Ultrastructural study of the blood cells of the beluga whale, Delphinapterus leucas.

General cytological and ultrastructural features of cells found in the peripheral blood of three captive beluga whales (Delphinapterus leucas) are described. We noted all major peripheral cell types common to the circulation of mammalian species, including polymorphonuclear neutrophils, eosinophils, basophils, lymphocytes, and monocytes, as well as myelocytes, erythroblasts, and plasma cells. Platelets and two populations (normal and hypochromic) of red blood corpuscles were observed.

Animals↗

Effects of alanine substitutions in alpha-helices of sperm whale myoglobin on protein stability.

The peptide backbones in folded native proteins contain distinctive secondary structures, alpha-helices, beta-sheets, and turns, with significant frequency. One question that arises in folding is how the stability of this secondary structure relates to that of the protein as a whole. To address this question, we substituted the alpha-helix-stabilizing alanine side chain at 16 selected sites in the sequence of sperm whale myoglobin, 12 at helical sites on the surface of the protein, and 4 at obviously internal sites. Substitution of alanine for bulky side chains at internal sites destabilizes the protein, as expected if packing interactions are disrupted. Alanine substitutions do not uniformly stabilize the protein, either in capping positions near the ends of helices or at mid-helical sites near the surface of myoglobin. When corrected for the extent of exposure of each side chain replaced by alanine at a mid-helix position, alanine replacement still has no clear effect in stabilizing the native structure. Thus linkage between the stabilization of secondary structure and tertiary structure in myoglobin cannot be demonstrated, probably because of the relatively small free energy differences between side chains in stabilizing isolated helix. By contrast, about 80% of the variance in free energy observed can be accounted for by the loss in buried surface area of the native residue substituted by alanine. The differential free energy of helix stabilization does not account for any additional variation.

Alanine↗

Phe-46(CD4) orients the distal histidine for hydrogen bonding to bound ligands in sperm whale myoglobin.

The role of Phe-46(CD4) in modulating the functional properties of sperm whale myoglobin was investigated by replacing this residue with Leu, Ile, Val, Ala, Trp, Tyr, and Glu. This highly conserved amino acid almost makes direct contact with the distal histidine and has been postulated to affect ligand binding. The overall association rate constants for CO, O2, and NO binding were little affected by decreasing the size of residue 46 step-wise from Phe to Leu to Val to Ala. In contrast, the rates of CO, O2, and NO dissociation increased 4-, 10-, and 25-fold, respectively, for the same series of mutants, causing large decreases in the affinity of myoglobin for all three diatomic gases. The rates of autooxidation at 37 degrees C, pH 7.0 increased dramatically from approximately 0.1-0.3 h-1 for wild-type, Tyr-46, and Trp-46 myoglobins to 1.5, 5.2, 4.9, and 5.0 h-1 for the Leu-46, Ile-46, Val-46 and Ala-46 mutants, respectively. Rates of NO and O2 geminate recombination were measured using 35 ps and 9 ns laser excitation pulses. Decreasing the size of residue 46 causes significant decreases in the extent of both picosecond and nanosecond rebinding processes. High resolution structures of Leu-46 and Val-46 metmyoglobins, Val-46 CO-myoglobin, and Val-46 deoxymyoglobin were determined by X-ray crystallography. When Phe-46 is replaced by Val, the loss of internal packing volume is compensated by (1) contraction of the CD corner toward the core of the protein, (2) movement of the E-helix toward the mutation site, (3) greater exposure of the distal pocket to intruding solvent molecules, and (4) large disorder in the position of the side chain of the distal histidine (His-64). In wild-type myoglobin, the van der Waals contact between C zeta of Phe-46 and C beta of His-64 appears to restrict rotation of the imidazole side chain. Insertion of Val at position 46 relieves this steric restriction, allowing the imidazole side chain to rotate about the C alpha - C beta bond toward the surface of the globin and about the C beta - C gamma bond toward the space previously occupied by the native Phe-46 side chain. This movement disrupts hydrogen bonding with bound ligands, causing significant decreases in affinity, and opens the distal pocket to solvent water molecules, causing marked increases in the rate of autooxidation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗