Vascular anatomy of the gills of the stingarees Urolophus mucosus and U. paucimaculatus (Urolophidae,Elasmobranchii)

The branchial vascular anatomy of Urolophus mucosus and U. paucimaculatus was studied by scanning electron microscopical examination of critical-point-dried tissue or of vascular corrosion casts. The vasculature could be divided into arterioarterial and arteriovenous pathways, which channel the flow of blood through the gills. The arterioarterial pathway consists of an afferent branchial artery which gives rise to afferent distributing arteries that run through the tissues of the interbranchial septum and supply the afferent filament arteries of several filaments. Afferent filament arteries open regularly into a corpus cavernosum in the core of the filament; unlike other elasmobranchs no septal corpora cavernosa are found. At the tip of the filament, channels of the corpus cavernosum connect to a channel which passes across the distal end of the filament from afferent to efferent side. This channel always connects to the afferent filament artery, and in many filaments it connects to the efferent filament artery as well. In addition, a vascular arcade connects all the afferent filament arteries along the entire length of each hemibranch. The filament corpus cavernosum supplies the secondary lamellae. The lamellae drain into efferent lamellar arterioles which in turn drain into the efferent filament artery and the efferent branchial artery. The vascular anatomy of the arteriovenous pathway is similar to that described in other elasmobranchs and consists of arteriovenous anastomoses, found only arising from efferent arterial circulation, and the venolymphatic system, which is composed of the central venous sinus and the companion vessels.     

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.     

Adrenergic innervation of the gills of brown and rainbow trout,Salmo trutta and S. gairdneri

The adrenergic innervation of structures in the gills of brown and rainbow trout was studied with catecholamine fluorescence histochemistry. In the arterio-arterial vascular pathway, there was an innervation of the afferent and efferent lamellar arterioles, but the afferent and efferent filamental arteries and the secondary lamellae were devoid of any fluorescent nerve fibres. In S. trutta only, there was an additional innervation of the afferent and efferent branchial arteries and the base of the efferent filamental artery. The innervation of the arterio-venous vascular pathway was similar in both trout species. Many fluorescent nerve fibres were found on nutritive arterioles in the gill arch and interbranchial septum, and in the core of each filament between the surface epithelium and the wall of the filament venous sinus. No fluorescent nerve fibres were observed at the origins of the capillaries arising from the efferent filamental artery. The sympathetic nerve supply is provided to the gills mainly through the posttrematic nerve, with an occasional small contribution through the pretrematic nerve. The presence of adrenergic nerves in the gills is discussed in relation to the regulation of blood flow through the arterio-arterial and arterio-venous pathways.     

Morphology and ultrastructure of the gills of terrestrial crabs (Crustacea,Gecarcinidae and Grapsidae): Adaptations for air-breathing

Summary The terrestrial crabsGeograpsus grayi, Geograpsus crinipes, Cardisoma hirtipes andGecarcoidea natalis have a reduced number of gills and show a reduced planar gill surface (SA) compared to aquatic species. Gill lamellae are stiffened and thickened (increasing blood/gas (BG) diffusion distances) and nodules maintain wide spacing between lamellae. Haemolymph is directed through the gill lamellae by rows of pillar cells and in the afferent region an intralamellar septum splits the haemolymph into two parallel networks. Gaps in the lines of pillar cells allow movement of haemolymph between adjacent channels. The afferent vessel distributes haemolymph to the lamella via a number of direct channels including the marginal canal and in large gills with the aid of a long, forked sinus which supplies the ventral and central regions of the lamellae. The marginal canal functions in both distribution and collection of haemolymph; the role varies with species. Potential flow-control sites were identified at the junctions between afferent and efferent areas and where the efferent channels enter the efferent branchial vessel. Each gill receives a branch from the sternal artery which supplies all the lamellae. Transport epithelia is the principal cell type in the gills of all species examined though its location varies between species, either being confined to certain gills or specific parts of the lamellae.The gill lamellae of air-breathing crabs are clearly modified to breathe air (stiffening and presence of nodules), though the overall contribution of the gills to gas exchange has been reduced (smaller SA and longer BG diffusion distances). The role of the gills in air-breathing crabs thus appears to have switched from one of an efficient aquatic gas-exchanger (thin with large surface area) and transport tissue, to one that is predominantly set up for ion-regulation.Abbreviations a afferent branchial vessel - ac afferent channels - art arteriole - ass artifactual subcuticular space - bl basal lamina - c cuticle - col collagen - ct connective tissue - e efferent branchial vessel - ec efferent channels - epi epithelium - f folds - g Glycogen - h haemolymph - hc haemocyte - is intralamellar septum - m marginal canal - mi mitochondria - mt microtubules - n nucleus - p pillar cell - s shaft of efferent vessel - sd septate desmosome     

The effect of input pressure and flow on the pattern and resistance to flow in the isolated perfused gill of a teleost fish

Summary When an isolated gill arch of the marine teleost,Ophiodon elongatus, was perfused under conditions which mimic those present in the intact animal, only two thirds of the gill lamellae were perfused. An increase in either input (afferent) pressure and flow or input pulse pressure caused an increase in the number of lamellae perfused as well as altering the distribution of the efferent outflow between the efferent artery and the venolymphatic drainage of the gill. The gill is compliant and an increase in efferent pressure reduced gill resistance to flow without altering the number of lamellae perfused. In these experiments there was no simple relationship between the number of lamellae perfused and gill resistance.These observations are of importance in the interpretation of results from pharmacological and ion exchange studies of isolated gills as well as indicating how cardiovascular changes could bring about alterations in gill blood flow in the intact fish.     

A morphological study on posterior gills of the mangrove crab Ucides cordatus

Morphological and histological studies on posterior gills of the mangrove crab Ucides cordatus showed that the 5th gill (of 7) has a larger surface area and a greater number of lamellae compared to the 6th gill. Regular separation of gill lamellae, important when the gill is in air, is maintained by enlargements of the marginal canals. Conical, spine-like structures along the efferent vessel of both 5th and 6th gills were also observed. In addition, pillar cells, a discontinuous lamellar septum and a hypobranchial artery were observed. The presence of valve-like structures near the efferent vessel was also indicated. These structures, together with the pillar cells, may have a role in directing the hemolymph flow towards certain gills during particular physiological states. Localization of osmoregulatory epithelia in the lamellae of both gills was inferred from dimethylaminostyrylethylpyridiniumiodine staining. Apparently gills 5 and 6 have osmoregulatory epithelial cell patches of similar area, corresponding to 43% and 38% of the total lamellae area, respectively. However, their localization is quite different. Gill number 5 osmoregulatory patches seem to be restricted to the afferent region of the lamella whereas in gill number 6, they are more dispersed over the entire lamella. These differences may be related to the particular functional characteristics of these gills.     

Structural elements of the gills of the shore crabCarcinus mediterraneus

Fine structural studies were conducted on the gills of the shore crabCarcinus mediterraneus using scanning electron microscopic techniques. The results obtained show the structural organization of crab gills from whole gills including spiny elements over the 150 lamellae to lamellar components such as cuticles, median shaft, marginal canal, afferent and efferent lamellar vessels and hemolymph cells. Enormous surface enlargement is accomplished by a variety of structural elements which allow rapid circulation of hemolymph. In the form of a relatively small organ, the gills fulfill all the necessary exchanges of specific molecules between the crab and its environment. Aggregations of ca 1-μm particles covering the outer cuticular surfaces are considered to be bacterial colonies of unknown properties and functions.     

Vascular anatomy of the fish gill

The fish gill is the most physiologically diversified vertebrate organ, and its vasculature the most intricate. Application of vascular corrosion techniques that couple high-fidelity resins, such as methyl methacrylate, with scanning electron microscopy yields three-dimensional replicas of the microcirculation that have fostered a better appreciate gill perfusion pathways. This is the focus of the present review. Three vascular networks can be identified within the gill filament. The arterioarterial (respiratory) pathway consists of the lamellae and afferent and efferent segments of the branchial and filamental arteries and lamellar arterioles. The body of the filament contains two post-lamellar pathways: the interlamellar and nutrient. The interlamellar system is an extensive ladder-like network of thin-walled, highly distensible vessels that traverses the filament between, and parallel to, the lamellae and continues around the afferent and efferent borders of the filament. Interlamellar vessels are supplied by short, narrow-bore feeder vessels from the medial wall of the efferent filamental artery. A myriad of narrow-bore, tortuous arterioles arise from the basal efferent filamental artery and efferent branchial artery and anastomose to form the nutrient circulation of the arch and filament. In the filament body, nutrient capillaries and interlamellar vessels are often closely associated, and the former may ultimately drain into the latter. Many of the anatomical characteristics of interlamellar vessels are strikingly similar to those of mammalian lymphatic capillaries, with the exception that interlamellar vessels are directly fed by arteriovenous-like anastomoses. It is likely that gill interlamellar and mammalian lymphatics are physiologically, if not embryologically, equivalent.     

An electron microscopic study on the innervation of the gill filaments of a lamprey,Lampetra japonica

This paper reports observations on the innervation of gill filaments of the lamprey, Lampetra japonica. Nerve fibers run on each side of the afferent filament artery (AFA nerve) and in the connective tissue compartment along the efferent filament artery (EFA nerve). The AFA nerve supplies vasomotor fibers to the afferent filament artery and arteriovenous anastomoses and special visceral motor fibers to branchial muscle fibers (musculus compressor branchialis circularis). Nerve endings of the vasomotor fibers contain large, cored vesicles (60–180 nm in diameter) with a variable number of small, clear vesicles (30–70 μm in diameter), whereas those of the visceral motor fibers have many small, clear vesicles with few large, cored vesicles. The EFA nerve supplies vasomotor fibers to the efferent filament artery. Their endings, containing mixtures of predominantly large, cored vesicles and small, clear vesicles make close synaptic contacts with reticular cells. The latter in turn are connected with each other or with smooth muscle cells in the wall of the efferent filament artery by nexuses. No nerves are found in the axial plate between the afferent and efferent filament arteries nor in the secondary lamellae of individual gill filaments. No afferent nerve supply to the gill filament has been found.     

Study of glomerular vasculature in teleosts with the resin-replica method

Glomerular vasculature was investigated in the carpCyprinus carpio, the scorpionfishSebastiscus marmoratus, and the marine catfishPlotosus lineatus with the resin-replica method. An afferent arteriole was connected with a glomerulus in every fish. It was slender in the carp, whereas the scorpionfish and marine catfish possessed thick afferent arterioles. The glomerular capillaries were sinusoidal. The divergences, convergences, and windings of these capillaries were not well developed in any of the fish. The glomerular capillaries converged into an efferent arteriole in the carp and scorpionfish. In the marine catfish, on the other hand, most of the glomeruli had two efferent arterioles.     

The morphology of gills of Haliotis tuberculata (Linnaeus, 1758)

Manganaro, M., Laurà, R., Guerrera, M.C., Lanteri, G., Zaccone, D. and Marino, F. 2011. The morphology of gills of Haliotis tuberculata (Linnaeus, 1758). —Acta Zoologica (Stockholm) 93 : 436–443. Although the morphology of abalone gills has been studied by some authors, up to date no data are available about the gills of Haliotis tuberculata. This study was carried out, by light and electron microscopy, on 10 wild adult H. tuberculata. Gills lamellae produce an undulated surface increasing the area in contact with water. At the level of skeletal rods, we observed a joint‐like structure that allows a checked movement. The left ctenidium is always decidedly larger than the right, probably because of the enormous size of the shell muscle. The cilia permit oxygenated water that leaves the afferent border and is thrust away at the tips of the lamellae by the extremely long cilia. Ciliary movement may take part in sweeping mucous secretions to capture extraneous particles and remove them from the gills. Three types of mucous cells are distributed along the epithelium of the afferent and efferent zones of the gill filament. They seem to play a role in the cleansing of gills in coordination with the muscle contraction and ciliary movement. The presence of microvilli on particular cells reflects their role associated with the absorption of substances from the environment. A haemolymphatic vessel is located in the central zone of the gill filament. The backbone of the haemolymphatic vessel is a chitino‐like structure, which gives support to the gills.     

Observations of valve-like structures and evidence for rectification of flow within the gill lamellae of the crab Carcinus maenas (Crustacea,Decapoda)

Summary The isolated gills of Carcinus maenas, perfused at pressure drops of 1–10 cm of water, exhibited flow rectification, the resistance to perfusion via the afferent vessel being many times lower than that for efferent perfusion. The asymmetry was greater at the lower end of this pressure range.The overall afferent branchial resistance for Carcinus of weight 65 g, and with no ventilatory component in the transmural pressure difference, was estimated to be 0.05 cm of water. l^–1 · sec. The corresponding overall reverse (efferent) branchial resistance was 0.36 cm of water · l^–1 · sec.LM, TEM and SEM examination of the gills indicated that haemolymph leaves each gill lamella via several discrete parallel efferent channels which drain different regions of the lamella, and that each efferent channel is nearly closed, at its junction with the efferent branchial vessel, by a diaphragm of loosely interwoven and very elongated cells. It is concluded that these cells may constitute efferent valves and that narrow apertures between them may contribute a major component to the branchial resistance and be primarily responsible for the rectification of flow. Relatively wide apertures lead directly from the afferent vessel into the lamellae and are not asociated with valves of any kind.The valves may be important in enabling changes in transmural pressure associated with ventilatory reversals to pump haemolymph unidirectionally through the lamellae. Similarly valves may allow the oscillating venous pressures associated with locomotor activity to improve gill perfusion during exercise.The elongated tails of the cells of the efferent valve contain numerous microtubules. The wider cell bodies contain the nucleus and many mitochondria. Unusual organelles composed of many short (about 0.25 m long) microtubules and often lying close to the nuclear membrane may be microtubule organising centres. It is speculated that, in addition to their simple mechanical function, the valve cells may play a more dynamic role in regulating flow of haemolymph through different lamellar routes, or that they may monitor composition, pressure or flow of the efferent lamellar circulation.     

Morphology and vascular anatomy of the gills of a primitive air-breathing fish,the bowfin (Amia calva)

Summary The morphology of the gills of a primitive air breather (Amia calva) was examined by light microscopy of semithin sections of gill filaments, and gill perfusion pathways were identified by scanning-electron microscopic analysis of corrosion replicas prepared by intravascular injection of methyl methacrylate. The arrangement of gill filaments and respiratory lamellae is similar to that of teleosts with the exception of an interfilamental support bar that is fused to the outer margins of lamellae on adjacent filaments. The prebranchial vasculature is also similar to that of teleosts, whereas the postbranchial circulation of arches III and IV is modified to permit selective perfusion of the air bladder. Gill filaments contain three distinct vascular systems: (1) the respiratory circulation which receives the entire cardiac output and perfuses the secondary lamellae; (2) a nutrient system that arises from the postlamellar circulation and perfuses filamental tissues; (3) a network of unknown function consisting of subepithelial sinusoids surrounding afferent and efferent margins of the filament and traversing the filament beneath the interlamellar epithelium. Prelamellar arteriovenous anastomoses (AVAs) are rare, postlamellar AVAs are common especially at the base of the filament where they form a dense network of small tortuous vessels before coalescing into a large filamental nutrient artery. Unlike in most teleosts, the outer vascular margins of the lamellae are embedded in the interfilamental support bar and become the sole vasculature of this tissue. Arterial-arterial lamellar bypass vessels were not observed. Previously observed decreases in oxygen transfer across the gills during air breathing can be explained only by redistribution of blood flow between or within the respiratory lamellae.Supported by NSF Grant No. PCM 79-23073The author wishes to thank Miss K. Drajus and D. Kullman for their excellent technical assistance and Dr. W. Gingerich, Mr. J. Crowther and D. Zurn for help in obtaining bowfin     

Vascular organization of the catfish gill filament

Summary Light and scanning electron microscopic observations were made on methyl-methacrylate corrosion casts of the blood vessels in the gills of channel catfish (Ictalurus punctatus). The vasculature of the gill filament can be divided into three distinct pathways: 1. the well-known respiratory circulation which includes the afferent filamental artery (AF), afferent lamellar arteriole (AL), lamella (L), efferent lamellar arteriole (EL) and efferent filamental artery (EF), 2. a nutritive pathway from the EF through small nutritive capillaries (NC) and into one of several filamental veins (FV), and 3. an interlamellar circulation in which small prelamellar arterio-venous anastomoses (PAVA) connect the AL into a series of organized vascular spaces (interlamellar vessels, ILV's) that underlie the interlamellar filamental epithelium. Several sinuslike spaces associated with AF, EF and the filamental cartilagenous support were also observed. The physiological significance of these vascular pathways is discussed.Supported in part by NSF Grant No. PCM 76-16840The authors wish to acknowledge the assistance of Mr. P. Holbert, Miss K. Drajus and Mrs. J. Smith. Gratitude is expressed by Kenneth R. Olson to Dr. Janice Nowell for her helpful suggestions with corrosion casting techniques     

白鲢鳃部血管的研究

本文用甲基丙烯酸甲脂铸型技术,配合扫描电镜,研究白鲢鳃部血管的分布。鳃区具有双血液循环,动脉—动脉循环,主司血液与水体间气体交换;动脉—静脉循环主司鳃区营养交换。入鳃丝动脉上具有膨大的“血泡”结构,可能起“鳃心”作用,当鳃丝内收时将血液输送入入鳃动脉。伪鳃鳃小片彼此排列紧密,这种结构似不利于气体交换,可能没有什么呼吸功能。     

The structure of the gill of the trout,Salmo gairdneri (Richardson)

Summary A light and electron microscopic study was made of the structure of the gill arch, filament and secondary lamella of Salmo gairdneri R. Blood pathways through the gill were traced from serial histological sections, and from the examination of ink perfused tissue and perspex casts formed following resin injection of the circulatory system.The epithelium covering the gill consists of unspecialized, dark, chloride and mucous cells. The distribution of specialized cells appears to be related to gill function. The basement membrane underlying the epithelium consists of three layers, the inner collagen layer being continuous with the connective tissue core of the gills.Blood supply to the secondary lamellar respiratory surface is via branchial, filament and secondary lamellar arteries. Blood spaces of the secondary lamellae are delimited by pillar cells containing what appears to be contractile material. The marginal channel of each lamella is bounded distally by cells of endothelial origin. A network of lymph spaces within the filaments connects with efferent branchial arteries. Nutritionary capillaries within the filaments connect with afferent branchial arteries. No shunts between afferent and efferent filament arteries were found.Data from this study and previous physiological and histopathological studies suggest a mechanism for the control of blood flow to suit the respiratory requirements of the fish. This mechanism involves a system of recruitment of additional respiratory units and changes in overall blood flow patterns.This work formed part of a thesis submitted for the degree of Doctor of Philosophy in 1971 and for which M. M. was in receipt of a studentship from the Natural Environmental Research Council. The authors are grateful for the support given by research grants from the M.R.C (P.T.) and the N.E.R.C. (M.M.), and to Prof. G. M. Hughes in whose department the work was carried out.     

波纹唇鱼鳃丝的光镜、扫描和透射电镜观察

应用光学显微镜、扫描电镜和透射电镜对波纹唇鱼(Cheilinus undulatus)鳃的组织结构、表面形态特征及鳃小片超微结构进行了观察.结果表明,波纹唇鱼有3对全鳃,1对半鳃和1对伪鳃,鳃丝呈梳状紧密排列在鳃弓上,鳃小片紧密地镶嵌排列在鳃丝两侧,入鳃动脉、出鳃动脉和鳃小片毛细血管网组成鳃的血液系统.鳃丝非呼吸区分布...     

The morphology of the gills of the freshwater African crab Potamon niloticus (Crustacea: Brachyura: Potamonidae): A scanning and transmission electron microscopic study

The gills of the African freshwater crab Potamon niloticus -Ortmann have been investigated by scanning and transmission electron microscopy. Potamon has seven pairs of phyllobranchiate gills contained in the branchial chambers. From the central axis of the gills arise bilaterally situated thin flaps, the lamellae. The afferent branchial vessel (the epibranchial vessel) is located on the dorsal aspect of the gill arch and the efferent vessel (the hypobrancial vessel) on the ventral side. Between these two blood vessels, the blood percolates through the lamellar vascular channels where it is oxygenated. The lamellae consist of an epithelial cell layer covered by a thin cuticle which consists of tightly fused but distinct layers. The epithelial cells approach each other at regular intervals and fuse in the middle of the lamellar sinus delineating the vascular channels. Apical profuse membranous infoldings and numerous mitochondria characterize the epithelial cells, features typical of cells involved in active transport of macro- and micromolecules. In Potamon , however, there were no distinct gas exchange and osmoregulatory regions of the gills. On average, the cuticle was 0.78 μm thick while the epithelial cell was 6 μm. Cells that were morphologically similar to the renal glomerular podocytes of the vertebrates were observed in the efferent gill vessel of Potamon. These cells have been said to be phagocytic and may play an important defensive role in the crustaceans. Although basically the morphology of the gills of Potamon is similar to that of the other decapods, fine structural differences were evident as would be intuitively expected in a group of animals that has undergone such remarkable adaptive radiation.     

Early development and microhabitat of the monogenean Horricauda rhinobatidis, with observations on the related Troglocephalus rhinobatidis, from Rhinobatos batillum from Queensland,Australia

The small monocotylid monogenean Horricauda rhinobatidis is abundant on the gills of its host Rhinobatos batillum, whereas the larger, related monogenean Troglocephalus rhinobatidis is comparatively uncommon. Young specimens of Horricauda live between the host's secondary gill lamellae. Post-oncomiracidia have 14 marginal booklets but as the larvae develop these are supplemented first by a pair of hamuli and then by muscular ventral loculi followed by six forwardly-directed, dorsal spines. By impaling secondary gill lamellae these spines may serve to prevent parasites from being dislodged by gill ventilating currents. Before reaching sexual maturity the parasites leave the secondary gill lamellae and establish themselves in the septal canals. It is uncommon to find more than one adult specimen of Horricauda in each septal canal. The significance of this in relation to sperm exchange is discussed. Like Troglocephalus, Horricauda has eight head sacs, and the suggestion is made that these sacs may play a part in feeding.     

The linear cable theory as a model of gill blood flow

The vascular organization of the teleost gill suggests that blood flow distribution from the filamental artery to the respiratory lamellae is governed by relationships analogous to the cable conduction properties of a nerve axon. The space constant (λ) by definition is the distance along the gill filament at which the in-series resistance of the afferent filament artery equals the in-parallel resistance of the afferent lamellar arteriolar, lamellar, efferent lamellar arteriolar (ALA-L-ELA) segments. Constriction of the afferent filamental artery or uniform dilation of the ALA-L-ELA will decrease λ. As λ decreases, flow through the proximal (basal) lamellae greatly increases at the expense of distal lamellar perfusion. When λ increases in a system of finite length the flow profile must account for reflected pressures within the main vessel. The λ calculated from corrosion casts of gill vasculature is $\text{1}\text{4}$ to $\text{1}\text{2}$ the filament length. This favors blood flow through the proximal lamellae and when cardiac output increases, the proportion of cardiac output perfusing the proximal areas increases at the expense of distal lamellar blood flow. To offset these changes it is proposed that increased distal lamellar perfusion is achieved by simultaneous vasodilatation of distal and constriction of proximal ALA-L-ELA segments and dilation of the afferent filamental artery.