Ultrastructural, Sensory and Functional Anatomy of the Northern Elephant Seal (Mirounga angustirostris) Facial Vibrissae.

Vibrissae (whiskers) play a key role in underwater orientation in foraging phocids through vibrotactile sensation processing. Our aim was to evaluate the structure of northern elephant seal (NES) vibrissae by means of light (LM) and transmission electron microscopy (TEM), in order to elucidate their function. Vibrissal follicles were processed using standardized laboratory methods and LM/TEM techniques. Individual follicular axonal numbers were counted and axonal diameter measured and averaged. NES have mystacial, rhinal, supraorbital and labial vibrissae. The vibrissal follicles are histologically subdivided into a ring, upper and lower cavernous sinuses (LCS). Each vibrissa is innervated by the deep vibrissal nerve. The average number of axons per large mystacial vibrissa is 1804 (±123), rhinal 985 (±241), supraorbital 1,064 (±204) and 374 (±65) in labial vibrissa. The entire vibrissal system carries an estimated 148 573 axons, and mystacial vibrissae alone have 125 323 axons. Axonal conduction velocity for each vibrissal type is 55.26 m/s for labial, 56.58 m/s for rhinal and 35.88 m/s for mystacial vibrissae. TEM and LM revealed a plethora of mechanoreceptors within the vibrissal follicles: Merkel cell-neurite complexes, lanceolate and pilo-Ruffini end organs. A vast number of sensory axons projecting from the entire vibrissal system indicate that the vibrissal sensory area takes up a large proportion of phocids' somatosensory cortex. In conclusion, NES has highly sensitive and finely tuned vibrotactile vibrissal sense organs.

Gross anatomy of the ringed seal (Pusa hispida) gastro-intestinal tract.

The gross anatomical structure of the ringed seal (Pusa hispida) gastrointestinal tract is poorly described and often veterinary anatomical terminology is not used. Although the basic abdominal visceral pattern corresponded to domestic carnivores, significant differences were noted. The stomach was an elongated sharply bent tube (u-shaped) with the pylorus and fundus juxtaposed. The elongated jejunum measured up to 15.6 times body length and had 37 jejunal arteries from the cranial mesenteric artery. The pancreas was asymmetrical with a small right lobe and a large left lobe. The unusually short greater omentum negated formation of deep and superficial leaves. The most remarkable difference was the separation of the liver parenchyma into three physically separate masses, held together by hepatic ducts, veins and arteries. The topography and position of the liver was dependent on the amount of blood in the hepatic sinus (distended hepatic veins and hepatic portion of vena cava). Thus, as the hepatic sinus filled, the lateral liver masses separate from the central mass by moving caudolaterally. This was facilitated by modified coronary and triangular ligaments which did not attach directly to the liver, but instead to the hepatic sinus. These anatomical adaptations are apparently advantageous to ringed seal's survival in a deep marine environment.

Histological assessment of selected blood vessels of the phocid seals (northern elephant and harbour seals).

Phocid seals exhibit vascular adaptations that allow them to undertake prolonged deep dives. These vascular adaptations are either unique to phocids, or are modified vascular equivalents to those present in terrestrial mammals. One such adaptation, the aortic bulb, is a spherical enlargement of the ascending aorta specific to phocid seals. Its histological make-up consists of a reinforced tunica media with circular and longitudinal layers of elastic fibres. This reinforcement enables multi-axial deformation of the aortic bulb, thus complementing its function as a prominent elastic reservoir or 'windkessel'. A second adaptation, the hepatic sinus, is an asymmetrical dilation of the abdominal portion of the caudal vena cava and accompanying hepatic veins. The hepatic sinus is comprised of a relatively thin tunica media, with a scant smooth muscle component. The bulk of the sinus wall is comprised of tunica adventitia. A third vascular adaptation distinctive to the phocids is the pericardial venous plexus, composed of convoluted veins circumnavigating the perimeter of the heart. Microscopically, these veins have a thick tunica media and also contain valves. Smaller arteries, venules and distinct capillary beds are observed interspersed in-between these veins. It can be hypothesized, that in seals, certain vascular embryonic development may be arrested at an earlier embryonic stage, resulting in these unusual vascular formations. These modifications play a vital role in blood pressure regulation and distribution of oxygenated blood during prolonged deep diving. The purpose of this work was to elucidate the histological aspects of these unique vascular modifications and relate them to specific function.

Macroscopic anatomy of the ringed seal [Pusa (Phoca) hispida] lower respiratory system.

This investigation serves to document the normal anatomical features of the lower respiratory tract of the ringed seal [Pusa (phoca) hispida]. Evaluation of embalmed specimens and tracheobronchial casts showed that the right lung of this seal consists of four lobes while the left has only three lobes. The ventral margins of the lungs do not reach the sternum causing them to form the boundary of the broad recessus costomediastinalis. Lung lobation corresponds with bronchial tree division. Pulmonary venous drainage includes right and left common veins draining ipsilateral cranial and middle lung lobes, and one common caudal vein draining both caudal lobes and the accessory lobe. The right and left pulmonary arteries divide into cranial and caudal branches at the level of the principal bronchus. The ringed seal has three tracheobronchial lymph nodes. The trachea has an average of 87 cartilages that exhibit a pattern of random anastomoses between adjacent rings. The trachea exhibits to a small degree the dorsoventrally flattened pattern that is described in other pinnipeds. The tracheal diameter is smaller than that of the canine.

Macroscopic anatomy of the great vessels and structures associated with the heart of the ringed seal (Pusa hispida).

The ringed seal [Pusa (Phoca) hispida], as well as other seals, exhibits unique anatomical properties when compared to its terrestrial counterparts. In the ringed seal, the most conspicuous marine adaptation is the aortic bulb. This large dilatation of the ascending aorta is comparable to that found in other seal species and marine mammals. The branches of the ascending aorta (brachiocephalic trunk, left common carotid artery and left subclavian artery) are similar to those of higher primates and man. The peculiarities of the venous system are: three pulmonary veins, a pericardial venous plexus, a caval sphincter, a hepatic sinus with paired caudal vena cavae and a large extradural venous plexus. Generally, three common pulmonary veins (right, left and caudal) empty into the left atrium. The pericardial venous plexus lies deep to the mediastinal pericardial pleura (pleura pericardica) on the auricular (ventral) surface of the heart. The caval sphincter surrounds the caudal vena cava as it passes through the diaphragm. Caudal to the diaphragm, the vena cava is dilated (the hepatic sinus), and near the cranial extremity of the kidneys, it becomes biphid. The azygos vein is formed from the union of the right and left azygos veins at the level of the 5th thoracic vertebra. Cardiovascular physiological studies show some of these anatomical variations, especially of the venous system and the ascending aorta, to be modifications for diving. This investigation documents the large blood vessels associated with the heart and related structures in the ringed seal.

Macroscopic anatomy of the heart of the ringed seal (Phoca hispida).

Anatomical properties of the ringed seal (Phoca hispida) heart and associated blood vessels reveal adaptations related to requirements for diving. Seven adult ringed seals were embalmed and dissected to document the gross anatomical features of the heart. Computed tomography images of the thoracic cavity were taken on one seal prior to dissection. The shape and position of the heart is different from the typical carnivore heart. The most notable difference is its dorsoventral flattened appearance with its right and left sides positioned, respectively, within the thoracic cavity. The long axis of the heart is positioned horizontally, parallel to the sternum. The right ventricle is spacious with thin walls which extend caudally to the apex of the heart such that the apex is comprised of both right and left ventricles. The cusps of the left atrioventricular valve of the ringed seal heart resemble an uninterrupted, circular curtain making it challenging to distinguish the divisions into parietal and septal cusps.

Microscopic anatomy of the ringed seal (Phoca hispida) lower respiratory tract.

The microscopic anatomy of the ringed seal lung exhibits unique features and many features similar to those described in other seal species. Unique features include: Trachealis muscle predominately oriented longitudinally; Large veins within the tracheal wall supported by elastic fibers; Goblet cells and pseudostratified epithelium lining the duct system of bronchial glands of the segmental bronchi; Lamina propria of the segmental bronchus heavily invested with elastic fibers clustered into dense longitudinal bundles; and Capillaries and venules covered with squamous epithelium protruding into bronchiolar lumina. Common features include: Cartilage support of the bronchial tree extending distally into respiratory bronchioles; Smooth muscle enhancements in the distal airways producing sphincter like formations; and Lungs extensively supported with interstitial tissue, which divide lungs into lobules.