The morphology and vasculature of the lungs and gills of the soldier crab,Mictyris longicarpus

The five gill pairs of Mictyris longicarpus have the lowest weight specific area reported for any crab. The cuticle of the gill lamellae is lined with epithelial cells which have structural features characteristic of iontransporting cells. Pillar cells are regularly distributed in the epithelium and serve to maintain separation of the two faces of the lamellae. The central hemolymph space is divided into two sheets by a fenestrated septum of connective tissue cells. The dorsal portion of the marginal canal of each lamella receives hemolymph from the afferent branchial vessel and distributes it to the lamella while the ventral portion of the canal collects hemolymph and returns it to the efferent branchial vessel. The lung is formed from the inner lining of the branchiostegite and an outgrowth of this, the epibranchial membrane. Surface area is increased by invagination of the lining which forms branching, blind-ending pores, giving the lung a spongy appearance. The cuticle lining the lung is thin and the underlyng epithelial cells are extremely attenuated, giving a total hemolymph/gas distance of 90–475 nm. Venous hemolymph is directed close to the gas exchange surface by specialised connective tissue cells and by thin strands of connective tissue which run parallel to the cuticle. Air sacs are anchored in position by paired pillar cells filled with microtubules. Afferent hemolymph is supplied from the eye sinus, dorsal sinus, and ventral sinus. Afferent vessels interdigitate closely with efferent vessels just beneath the respiratory membrane. The two systems are connected by a “perpendicular system” which ramifies between the airways and emerges to form a sinus beneath the carapace and then flows back between the air sacs to the efferent vessels. The afferent side of the perpendicular system is the major site of gas exchange. Efferent vessels return via large pulmonary veins to the pericardial cavity. P_aO₂ levels were high (95.5 Torr), indicating highly efficient gas exchange.     

Land crabs with smooth lungs: Grapsidae,Gecarcinidae, and Sundathelphusidae ultrastructure and vasculature

The highly terrestrial grapsids and gecarcinids and the amphibious sundathelphusids all have large, expanded branchial chambers. The lining of the branchial chambers is smooth and well vascularized, and it functions as a lung. The respiratory membrane and the cuticle lining the lung are extremely thin (200–350 nm). The blood vessels within the lung are formed from connective tissue cells supported by collagen fibres and lined by a basal lamina. The major vessels in the lung are embedded deep in the branchiostegite and lie just beneath the thick outer carapace. These vessels branch towards the respiratory membrane, where they eventually lose their connective tissue coverings to form thin, flattened lacunae directly below the respiratory epithelium. The lacunae (exchange sites) are bordered by specialized connective tissue cells, which either bear microvilli on their apical surface (fimbriated cells) or are very smooth. The respiratory circulation in the lung is very complex, with two portal systems present between the afferent and efferent systems, producing a total of three lacunal exchange beds. Portal systems increase the surface area available for gas exchange. The major distributing vessel in the lung is the branchiostegal vein, which runs along the inner margin of the branchiostegite. The main venous supplies come anteriorly from the infraorbital and ventral sinuses and posteriorly from the procardial sinus. The main collecting vessel is the pulmonary vein, which arises anteriorly and which runs around the ventral perimeter of the branchiostegite before emptying into the pericardial sinus. © 1993 Wiley-Liss, Inc.     

The morphology and vasculature of the respiratory organs of terrestrial hermit crabs (Coenobita and Birgus): gills,branchiostegal lungs and abdominal lungs

The morphology and vasculature of the respiratory organs of the terrestrial coenobitids were studied using light microscopy, TEM, SEM and corrosion casting. The gills of Coenobita and Birgus are modified for air-breathing but are reduced in number and size and have a comparatively small surface area. The branchiostegal lungs of Coenobita (which live in gastropod shells) are very small but are well vascularized and have a thin blood/gas barrier. Coenobita has developed a third respiratory organ, the abdominal lung, that is formed from highly vascularized patches of very thin and intensely-folded dorsal integument. Oxygenated blood from this respiratory surface is returned to the pericardial sinus via the gills (in parallel to the branchiostegal circulation). Birgus, which does not inhabit a gastropod shell, has developed a highly complex branchiostegal lung that is expanded laterally and evaginated to increase surface area. The blood/gas diffusion distance is short and oxygenated blood is returned directly to the pericardium via pulmonary veins. We conclude that the presence of a protective mollusc shell in the terrestrial hermit crabs has favoured the evolution of an abdominal lung and in its absence a branchiostegal lung has been developed.     

Structure of sinuses in the human lymph node

Summary A casting technique has been employed to display in three dimensions, the lymphatic microcirculation within the human lymph node. The casting compound filled the marginal sinus, and diffusely permeated the cortical lymphoid parenchyma. However, deep within the lymph node in the medullary region, the medium remained within the limits of the sinus walls. The casts showed well-defined channels appearing similar to vessels. These converged into larger vessels, which drained into efferent lymphatics leaving the node at the hilus.Electron microscopic examination showed that the outer wall of the marginal sinus and the trabecular side of trabecular sinuses had an intact, continuous endothelium with a basement membrane. However, gaps were present in the inner wall of the marginal sinus, as well as in the parenchymal wall of the trabecular sinus. In the medulla, the sinuses were lined by endothelial cells which appeared similar to macrophages. The sinus lining was incomplete and possessed numerous perforations. These observations indicated that sinus walls adjacent to connective tissue served as a barrier to cell movement, but those adjacent to a large lymphoid cell population had gaps, with cells in apparent transit between sinus lumen and parenchyma.     

Vascular pathways in the gill filaments of the flounder, Platichthys flesus L.

This study is concerned with functional organization of some of the blood pathways in the gill filament of the flounder, Platichthys flesus L. The existence of two independent vascular pathways has been confirmed. The blood from the efferent filament artery (EFA) enters the central venous sinus (CVS) through very small blood vessels which are characterized by the presence of sphincter-like structures. The existence of an independent chamber of the CVS mainly full of white blood cells provides evidence of an independent lymphatic system connected to the CVS. Gill rays support the afferent side of a gill filament whereas plasma and an extensive network of nutritive blood vessels in the CVS supports the efferent part.     

Histology of the caprine hemal node

Caprine hemal nodes were studied by light microscopy after glutaraldehyde fixation and epoxy resin embedding. A node consisted of a capsule, subcapsular and other sinuses, cortex, medulla and hilus. Elements of circulating blood filled the interstices of the reticular meshwork and associated macrophages which traversed the lumina of subcapsular and medullary sinuses. The latter were rare in 1-month-old goats, progressively increased in number and size in 2- to 4-month-old goats and coalesced with each other and the subcapsular sinus in adult animals. The cortical tissue appeared as lymphoid nodules. Circumferential lymphatic vessels abutted on outer margins of the nodules and gave origin to several radial lymphatics which branched and anastomosed between the medullary blood sinuses. Medullary cords were organized around the radial lymphatics. A single efferent lymphatic was formed at the hilum by confluence of the radial lymphatics. Our study, in contrast to earlier reports, shows that caprine hemal nodes possess efferent lymphatics. The present data suggest that the hemal nodes are involved, in addition to classical functions, in blood storage by hemoconcentration.     

Arterial system of the gill of the lobster,Homarus americanus

Three-dimensional architecture of the branchial artery and venous vasculature of Homarus americanus was studied by the method of corrosion cast or styrene cracking and by scanning electron microscopy. Four arteries, the epibranchial (EA) and hypobranchial arteries (HA) on the septal wall of the afferent and efferent vessels, respectively, and two lateral canal arteries (LCA), each in one of the paired lateral canals, run parallel to the gill axis. The EA directs dendroid branches to the spongy tissue in the afferent vessel wall far from the efferent, supplying oxygen to the otherwise oxygen-depleted tissue. The HA distributes the filament arteriole (FA) into the central channel of individual middle filaments via the LCA. The FA opens halfway at a position where the channel narrows. Thus, it is likely that venous hemolymph in the central channel flows from base to tip in the direction in which arterial hemolymph from the FA flows. This and the anatomy of venous vasculature suggest three probable patterns of perfusion from afferent to efferent vessels: double serial circulation via the outer and inner filaments and novel routes both through the middle filament, i.e., single circulation via the afferent and efferent channels of this filament and double serial circulation via the outer filament and then the central channel of the middle. On the basis of the physics of flow and known physiological data, we propose that switching of these routes that involves independently functional multiple double serial circulations can play an important role in controlling efficiency of gas exchange, particularly during hypoxia. J Morphol. 233:165–181, 1997. © 1997 Wiley-Liss, Inc.     

Renal vascular anatomy of the toad (Bufo marinus)

The renal vasculature of the toad, Bufo marinus, was studied mainly by means of scanning electron microscopy of vascular corrosion casts. All arterial branches terminated in a glomerulus. Each glomerulus was supplied by only one afferent arteriole. No shunts between afferent and efferent arterioles were observed. The glomerular channels appeared to be permanent capillaries. No evidence supporting the theory of freely shifting glomerular blood channels was found. Efferent arterioles radiated out towards the dorsal surface of the kidney where they connected with peritubular vessels. The renal portal veins produced an anastomosing plexus on the dorsal surface of the kidney, giving rise to the peritubular vessels. Peritubular vessels ran radially toward the ventral surface of the kidney, where they formed the roots of the renal veins. Attention is drawn to the possibility of hairpin countercurrent exchange between the capillary-like efferent arterioles and the peritubular vessels in the dorsal kidney.     

Accessory hemolymph pump in the mesothoracic legs of locusts, (Schistocerca gregaria forskal) (Orthoptera,Acrididae)

An accessory pulsatile organ located in the mesothoracic legs pumps hemolymph towards the tip of the leg ventrally and towards the body near the dorsal side. It consists of a muscle attached to the ventral side of the trochanter and to the central region of a transverse connective tissue diaphragm located at the trochanter-femur border. The diaphragm has a ventral outlet that permits efferent hemolymph flow through a narrow femoral sinus. A second dorsal outlet allows the afferent countercurrent back to the thorax through a separate hemolymph channel. During abdominal ventilation, the pumping rhythm of the legfn2heart is neurally synchronized with abdominal ventilation. Expiratory pressure expands tracheal air sacs in the ventral trochanter and helps driving hemolymph out of this space. In idle periods of resting ventilation, an autonomous myogenic rhythm of the leg–heart can maintain hemolymph circulation in the mesothoracic leg without neural control.     

Ultrastructure of the lung of the rattlesnake,Crotalus viridis oreganus

Scanning and transmission electron microscopic observations were made on the rattlesnake lung, which has the form of a cigar-shaped bag enclosing a large axial air chamber. The lungs were fixed by tracheal instillation of fixative to preserve the structural features of inflated lungs. An open tracheal groove along the ventral aspect of the lung is the only structural “airway” present. The wall of the lung has two histologically distinct regions: anteriorly, a respiratory portion, where up to three generations of septa subdivide the wall into cup-shaped gas-exchange chambers, termed faveoli; and posteriorly, a simple, thin-walled saccular portion. The epithelium lining the internal surface of the lung is composed of several cell types: (1) ciliated cells; (2) type I pneumonocytes; (3) type II pneumonocytes, secretory cells characterized by the presence of lamellar bodies; and (4) serous epithelial cells, secretory cells characterized by the presence of homogeneous, densely staining secretory granules. However, the distinctiveness of the secretory cell types in the snake lung is blurred because intermediate-appearing cells have both the lamellar body and homogenous type of secretory granule. The nonepithelial components of the pulmonary wall and septa consist of blood vessels and lymphatics, smooth muscle cells and fibroblasts, embedded in a matrix of extracellular connective tissue fibers. Tubular myelin figures were observed in the faveolar lining layer.     

Book lung development in the embryo,postembryo and first instar of the cobweb spider,Parasteatoda tepidariorum C. L Koch, 1841 (Araneomorphae,Theridiidae)

Light and electron microscopy were used to compare spider book lung development with earlier studies of the development of horseshoe crab book gills and scorpion book lungs. Histological studies at the beginning of the 20th century provided evidence that spider and scorpion book lungs begin with outgrowth of a few primary lamellae (respiratory furrows, saccules) from the posterior surface of opisthosomal limb buds, reminiscent of the formation of book gills in the horseshoe crab. In spider embryos, light micrographs herein also show small primary lamellae formed at the posterior surface of opisthosomal limb buds. Later, more prominent primary lamellae extend into each book lung sinus from the inner wall of the book lung operculum formed from the limb bud. It appears most primary lamellae continue developing and become part of later book lungs, but there is variation in the rate and sequence of development. Electron micrographs show the process of air channel formation from parallel rows of precursor cells: mode I (cord hollowing), release of secretory vesicles into the extracellular space and mode II (cell hollowing), alignment and fusion of intracellular vesicles. Cell death (cavitation) is much less common but occurs in some places. Results herein support the early 20th century hypotheses that 1) book lungs are derived from book gills and 2) book lungs are an early step in the evolution of spider tracheae.     

Functional Organization of the Teleost Gill

The circulation of the gills has been studied in the perch, trout and eel combining the conventional histological methods and casting techniques. The existence of two blood pathways in each gill arch was confirmed. 1 — An arterio-arterial pathway assuming the respiratory function. It includes the afferent branchial artery and in each primary lamella the afferent primary artery, the secondary lamellae capillaries and the primary and branchial efferent arteries. 2 — An arterio-venous pathway arising from both the branchial artery, in the gill arch, and the primary arteries in each primary lamella. This pathway includes the central venous sinus of the primary lamella, several small veins and is finally connected with the branchial veins. 3 — The lack of connections between afferent primary arteries and cvs in the trout and the perch makes impossible a direct blood flow from the afferent to the efferent artery (shunt). In the eel connections between cvs and both afferent and efferent arteries do not mean that a shunt is operating according to the pressure gradient.     

Light and electron microscopic study of the hindgut of the ant (Formica nigricans, hymenoptera): II. Structure of the rectum

The rectum of the ant Formica nigricans is composed of six ovoid rectal papillae inserted into a rectal pouch. The wall of the rectal pouch is made up of a flat epithelium of simple rectal cells lined by cuticle, and surrounded by a circular muscle layer. Each rectal papilla is comprised by a simple columnar epithelium of principal cells facing the lumen, and a simple cuboid epithelium of secondary cells towards the hemolymph; a group of 20-25 slender junctional cells lies laterally between both epithelia enclosing an intrapapillar sinus. The muscle layer of the rectal wall also surrounds the base of the papillae. Principal cells do not exhibit extensive infoldings at the apical and basal plasma membranes. Lateral membranes, in contrast, develop highly folded mitochondria-scalariform junction complexes enclosing very narrow intercellular canaliculi between adjacent cells. These canaliculi open to wider intercellular sinuses that ultimately drain into the intrapapillar sinus at the sites of entry of tracheal cells. The lateral plasma membranes do not link to the apical or basal plasma membrane, thus originating a syncytium throughout the principal cells. The apical plasma membrane of secondary cells shows invaginations in relation with an apical tubulovacuolar system, bearing portasomes to the cytoplasmic side of the membrane. Secondary cells unite by convoluted septate junctions, and basolateral infoldings are also developed. These ultrastructural traits, some of them different from those found in other insects, are discussed and examined in relation to their role in water and solute absorption. A route for rectal transport in F. nigricans is proposed.     

Aspects of the circulatory system of Pollicipes polymerus J. B. Sowerby (Cirripedia: Thoracica)

The circulatory system of Pollicipes polymerus exhibits a high degree of organization which precludes it from being referred to as an open system. The system is arbitrarily divided into four parts: (1) the circulation of the peduncle and mantle; (2) the distributive circulation of the body, which provides hemolymph to most of the cephalic gut, to the maxillary gland, and to the cirri; (3) the peripheral circulation which distributes blood from the cirri to the peripheral areas of the thoracic region, to most of the thoracic gut, and from the scutal sinus to the peripheral areas of the cephalic region; and (4) the collecting circulation, which conveys hemolymph mostly from the peripheral circulation of the body to the peduncle. There also may be a circulation that is comparable to the vertebrate lymphatic system. Pumping of hemolymph can be attributed to three pairs of skeletal muscles that compress the dorsolateral channels. These muscles are unique for crustacean muscles in that they do not appear to be striated. The rostral vessel appears to be a vestige of a heart in which the pump muscles have been lost. There is a similarity of the rostral vessel to the heart of Calanus finmarchicus (a copepod). This is additional evidence linking the cirripeds with the copepods within the Maxillopoda. Electron microscope observations of the walls of the midsagittal vessels indicate that there is a more or less random layering of cellular and noncellular elements within the wall. Muscle cells appear to be incorporated in the vessel wall.     

Effects of paranasal sinus ostia and volume on acoustic rhinometry measurements: a model study.

We used pipe models to investigate the effects of paranasal sinus ostium size and paranasal sinus volume on the area-distance curves derived by acoustic rhinometry (AR). Each model had a Helmholtz resonator or a short neck as a side branch that simulated the paranasal sinus and sinus ostium. The AR-derived cross-sectional areas posterior to the ostium were significantly overestimated. Sinus volume affected the AR measurements only when the sinus was connected via a relatively large ostium. The experimental area-distance curve posterior to the side branch showed pronounced oscillations in association with low-frequency acoustic resonances in this distal part of the pipe. The experimental results are discussed in terms of theoretically calculated "sound-power reflection coefficients" for the pipe models used. The results indicate that the effects of paranasal sinuses and low-frequency acoustic resonances in the posterior part of the nasal cavity are not accounted for in the current AR algorithms. AR does not provide reliable information about sinus ostium size, sinus volume, or cross-sectional area in the distal parts of nasal cavity.     

Homology and evolution of the orbitotemporal venous sinuses of humans.

The orbitotemporal venous sinuses accompany the intracranial branches of the stapedial artery. These sinuses are large in primitive primates and drain the extensive territories supplied by the stapedial artery as well as the brain. The orbit is drained by a wide cranio-orbital sinus which empties into the postglenoid emissary vein. Also emptying into the postglenoid vein is the petrosquamous sinus. The latter diverts cerebral blood from the transverse sinus and also drains the temporalis muscle. Emptying into both the cranio-orbital and petrosquamous sinuses are meningeal tributaries, which drain the cranial side wall and the dura mater. The relatively small sinus communicans runs in the angle between the petrosal bone and the cranial side wall. It commences at the postglenoid vein and connects the distal end of the petrosquamous sinus to the pterygoid venous plexus. In humans, the orbitotemporal sinus system is greatly modified. Its remnants persist for the most part as "middle meningeal veins." The system no longer drains the orbit, the temporal fossa, or the brain. The petrosquamous sinus becomes attenuated or obliterated along part or all of its length. The postglenoid vein vanishes. The cranio-orbital sinus is reduced in diameter and its connection to the orbit is feeble or absent. During development, the posterior end of the cranio-orbital sinus migrates inferiorly along the sinus communicans. In most individuals, this migration ceases at the foramen spinosum, site of the emissary vein of the sinus communicans. Meningeal tributaries are relatively large in humans, and drain principally into the cranio-orbital sinus or sphenoparietal sinus. The sphenoparietal sinus is an evolutionary novelty restricted to hominoids and is frequently developed in only Homo and Pongo.     

A morphological study on gills of the brown shrimp, Penaeus aztecus

The gills of Penaeus aztecus were examined by light and electron microscopy. They are dendrobranchiate, consisting of a central axis with biserially arranged branches that subdivide into bifurcating filaments. A septum divides the lumina of these structures into afferent and efferent channels. Hemolymph from the sternal sinus flows through the afferent channels into the filaments where it is directed into the efferent channels and finally to the pericardial cavity. In addition to these channels, numerous blood vessels permeate the gill. The cuticle covering the gill overlies a thin epithelium which is separated from hemolymph by a basal lamina. The epithelium, which is active in cuticle secretion, has a series of pillar processes that form subcuticular lacunae. The apical membranes of epithelial cells become folded in shrimp exposed to hypo- and hyperosmotic salinities. Granular cells that contain elaborate Golgi apparati and several types of granules are present throughout the gill. Nephrocytes resembling glomerular podocytes line the efferent channels. A large nerve traverses the septum in the axis.     

Die Vaskularisation der Kiemen von Myxine glutinosa L. (Cyclostomata)

The angioarchitecture of the gills in Myxine glutinosa L. was studied by scanning electron microscopy of vascular corrosion casts (methylmethacrylate). It was found that the afferent branchial artery may be connected to the sinus peribranchialis by a papilla. The sinus is connected to delicate vessels and sinuses, which are interposed between meridionally arranged radial arteries. These delicate vascular formations continue into the interior of the gill and form a plexus on both poles of the gill folds. Light and scanning electron microscopical studies on the vascular endothelium of the afferent vessels of the gill lamellae reveal rounded endothelial cells, which are characterized by a high content of granula, by long cellular processes and by bridge-formations towards neighbouring cells. Supporting columns within the vascular system of the lamellae were found to be set up by several spirally arranged cells. SEM observations reveal goblet cells and numerous superficial epithelial cells with various numbers of microvilli and microridges forming the epithelial surface of the gill folds.     

Vascular anatomy of the gills in a high energy demand teleost,the skipjack tuna (Katsuwonus pelamis)

Tunas (family: Scombridae, Tribe: Thunnini) exhibit anatomical, physiological, and biochemical adaptations that dramatically increase the ability of their cardiorespiratory systems to transfer oxygen from the water to the tissues. In the present study the vascular anatomy of the skipjack tuna, Katsuwonus pelamis, gill was examined by light and scanning electron microscopic analysis of methyl methacrylate vascular corrosion replicas prepared under physiological pressure. The gill filament contains three distinct blood pathways, respiratory, interlamellar, and nutrient. The respiratory, or arterio-arterial (AA) pathway, is the site of gas exchange and consists of the afferent and efferent filamental arteries (AFA and EFA) and arterioles (ALA and ELA) and the lamellae. Each ALA in the basal filament supplies ten or more lamellae and they anastomose with their neighbor to form a continuous vascular arcade. Four modifications in the lamellar circulation appear to enhance gas exchange efficiency. 1) The ALA deliver blood directly to the outer margin of the lamellae where unstirred boundary layer effects are predicted to be minimal and water PO2 highest. 2) Pillar cells are closely aligned along the outer boundary of the inlet side and the inner boundary of the outlet side of the lamellae to form multiple distributing and receiving blood channels. 3) Elsewhere in the lamella, pillar cells are aligned to form diagonal channels that direct blood from the outer to the inner lamellar margins, thereby reducing vascular resistance. 4) The lamellar sinusoid is especially widened near the efferent end to augment oxygen saturation of blood flowing through the inner margin. These adaptations, plus the presence of a bow-shaped interlamellar septum, and a thinned filament core appear to decrease gill vascular resistance and maximize gas-exchange efficiency. The interlamellar (IL) and nutrient systems originate from post-lamellar vessels and are arterio-venous (AV) pathways. IL vessels form an extensive ladder-like lattice on both sides of the filamental cartilage and are supplied in part by narrow-bore vessels from the medial wall of the EFA. Their function is unknown. Nutrient vessels are formed from the confluence of a myriad of tortuous, narrow-bore vessels arising from the basal region of the EFA and from efferent branchial arteries. They re-enter the filament and eventually drain into the IL system or filamental veins. As these AV pathways are retained despite considerable reduction in filamental tissue, it is evident that they are integral components of other non-respiratory homeostatic activities of the gill.