Macrophages and Reticulum Cells in the Spleen of the Dogfish,Scyliorhinus canicula

The presence and ultrastructural features of reticulum cells and macrophages were studied in the spleen of the dogfish Scyliorhinus canicula. Three morphologically distinguishable regions of the spleen were identified: the white pulp, the red pulp and the ellipsoids. In all three, the splenic parenchyma was a meshwork supported by reticulum cells and fibres. Reticulum cells in both the white and the red pulp are irregular elements, the processes of which are joined by cell junctions and embrace developing reticular fibres. The ellipsoids of the dogfish spleen are terminal branches of the splenic arteries of the white pulp, with a sheath consisting of reticulum cells, reticular fibres, ground substance, macrophages and occasional lymphocytes. Isolated melanomacrophages also occur in the ellipsoid walls as well as in the red pulp. In both the white and the red pulp phagocytic reticulum cells, and macrophages appear frequently forming cell associations with surrounding blood cells, mainly lymphocytes. The functional significance of the ellipsoids and the cell-cell clusters of the white and the red pulp is discussed in relation to the immune capacities demonstrated in elasmobranchs.     

Ultrastructure of splenic white pulp of the turtle,Mauremys caspica

Summary The ultrastructure of splenic tissue of non-immunized turtles, Mauremys caspica, shows two areas, namely, the white pulp which is lymphoid in nature, and the red pulp which is formed by cell cords and sinusoids. Between both areas there is always a marginal zone with gaps through which cells leak. In the white pulp, there are two blood vessel types; one with muscled walls, and the other showing thinner walls sheathed by reticular cells. Reticular cells constitute a network where there occur dendritic macrophages, lymphoblasts and small and medium lymphocytes. Mature plasma cells are scarce in the white pulp.     

Structure and histochemical organization of the spleen of Agama stellio (Sauria: Agamidae) and Chalcides ocellatus (Sauria: Scincidae)

The spleen of Agama stellio is composed mainly of red pulp; the white pulp is poorly developed, and its clusters are scattered throughout the organ and contain lymphocytes, reticular cells, and some plasma cells. The red pulp consists of clear reticular cells intermingled with blood cells, sinusoids, and pigment cells. The spleen of Chalcides ocellatus is encapsulated by connective tissue and is composed of white and red pulp. The white pulp consists of lymphoid tissue that surrounds the central arterioles, forming the periarteriolar lymphocyte sheath (PALS). The red pulp is composed of a system of venous sinuses and cords. The results of various histochemical procedures designed to demonstrate mucosubstances, proteins, and nucleic acids indicate that the spleen in these species resembles the mammalian spleen. © 1993 Wiley-Liss, Inc.     

Ultrastructure of phagocytic cells in the spleen of Psammophis sibilans (serpentes: Colubridae)

The spleen of Psammophis sibilans is composed mainly of red pulp, the white pulp being poorly developed. The white pulp lymphoid clusters are scattered throughout the organ and contain lymphocytes, reticular cells, and some plasma cells. The red pulp consists of reticular cells intermingled with blood cells, sinusoids, and melanomacrophage centers (MMCs). Filtering of particulate matter from the blood occurs in the red pulp by phagocytes of the pulp cord. MMCs are formed by the association of free macrophages that have phagocytosed some blood cells. Early filtering of particulate matter by the phagocytes of the pulp cords may allow for more efficient phagocytosis of erythrocytes by the MMCs. © 1994 Wiley-Liss, Inc.     

The argentophil reticular cells of the mammalian spleen. II. The argentophil reticular cell arrangement inside the red pulp and its relationship with the endothelial lining cells of the splenic sinuses

The red pulp's argentophil reticular cell network of the spleen is composed by 3 types of fixed cells: 1. the primitive reticular cell, slightly argentophil; 2. the small reticular cell; 3. the larger reticular cell, strongly argentophil and phagocytic. This latter shows the classical morphological characteristics attributed to the reticular cells of the spleen. The large argentophil reticular cell may become free, constituting a 4th cell type, the free macrophage. A 5th reticular cell type is the dendritic cell found into the lymphatic follicles of the white pulp. The argentophil reticular cells of the red pulp assemble together to form the reticular cells' network, that occurs inside the red pulp cords. The primitive and the small reticular cell form the fundamental network on which the large cells are apposed. The reticular cells of this network maitain relationship with the arterial terminal vessels of the red pulp, being responsible by the ellipsoid structure. In those arteriolar segments without ellipsoid and in those mammalian species devoid of ellipsoid, the white pulp reticular cells, that surround the blood vessel as a part of the lymphoid periarteriolar sheath, mix with the red pulp's reticular cells and both can hardly be discriminated. The ellipsoids are formed by large argentophil cells arranged in concentrical layers around its lumen that sometimes appear devoid of endothelial lining cells. The red pulp's argentophil reticular cells, either the small or the large ones, contributed to the structure of the splenic sinuses' wall; its thin processes surround the sinus wall outside the endothelial lining cell as fibrillar structures that cross the back side of the lining cells. Two or more argentophil reticular cells send fibrillar processes to a single sinus. The perisinusal reticular cells may send a process between adjacent endothelial lining, cells that insinuate and attain the sinus lumen; this process becomes thick and eventually, the reticular cell enter the sinus lumen as a free macrophage. The argentophil reticular cells of the red pulp make connection between the capsule or the trabeculae and the reticular cell network. The endothelial lining cells of the splenic sinuses are not argentophil.     

The intermediate circulation in the nonsinusal spleen of the cat, studied by scanning electron microscopy of microcorrosion casts

Scanning electron microscopy of microcorrosion casts was used to visualize circulatory pathways of the intermediate circulation in nonsinusal spleen of cat. The marginal sinus (MS) around lymphatic nodules is a distinct vascular space which fills preferentially before the filling of the marginal zone (MZ) and surrounding red pulp occurs. The MS, which has a plentiful vascular supply, does not usually enclose the nodule completely. From the MS, flow occurs radially outwards into the MZ. Corrosion casts and histological sections both showed that a diversity of forms of the MZ exists: The thickness of MZ and the arrangement of its reticulum vary among nodules and between different areas of the same nodule, from a complete absence to a region of up to 50 microns in width. No direct arteriovenous connections were found (in contrast to dog spleen: Schmidt et al., '83b). Aside from capillary endings in the MS and MZ, all arterial capillaries terminate in the reticular spaces of the red pulp, i.e., the circulation appears to be entirely "open." From each capillary termination a great variety of flow pathways through the reticular meshwork to the pulp venules is available; some of these routes are quite long but others may involve distances as short as 15-25 microns. Evidence of flow into ellipsoid sheaths was abundant in casts from dilated spleens, but scarce in contracted spleens. In contrast to the extensive system of interconnected venous sinuses in dog spleen, the pulp venules found in cat spleen are nonanastomosing, shorter, and much smaller in caliber, and all receive flow freely from the reticular meshwork via open ends and fenestrations in their walls.     

Structure of the spleen of the sea bass (Dicentrarchus labrax): A light and electron microscopic study

The spleen of sea bass (Dicentrarchus labrax) is composed mainly of red pulp, whereas the white pulp is poorly developed. The red pulp consists of clear reticular cells intermingled with blood cells, sinusoids, and melanomacrophage centers (MMCs). The MMCs are enclosed by an interrupted connective tissue capsule and show some areas in continuity with the adjacent pulp. The MMCs are formed by the association of free macrophages that have phagocytosed some blood cells. Sparse white pulp is diffuse, forming a cuff around the pulp arteries and MMCs, or occurring in small groups between the splenic cords. A longitudinal artery and vein, lying side by side, extend the length of the spleen. Frequently the capillaries are surrounded by a sheath of macrophages or ellipsoids. These macrophages may contain erythrocytes in varying degrees of degradation. Lymphopoiesis and plasmapoiesis occur in the sparse lymphold areas. Abundant plasma cell groups may indicate the presence of antibody production.     

鲤鱼和鲫鱼脾脏显微结构和亚显微结构的研究

鲤鱼和鲫鱼的脾脏中都没有小梁,其红髓和白髓是混合的。脾脏中都有椭圆体,其末端向脾髓开放。脾髓的纤维网眼中缺乏血管壁,椭圆体毛细血管中的血液可直接流入脾髓网状结缔组织的腔隙中。腹腔注射墨汁的鲫鱼,2小时后即可在椭圆体的巨噬细胞中看到碳粒的积累,可见脾脏的过滤作用主要在椭圆体中进行。在注射墨汁的鲫鱼脾脏中,黑色素巨噬细胞中心比未经注射鱼的脾脏明显,且数量增多,它和哺乳动物淋巴结和脾脏中的生发中心结构和功能都不相同。       

Phagocytosis in the spleen of the sunfish Lepomis spp.

A histological investigation of the filtering function of the spleen of the sunfish Lepomis spp. was conducted by light, scanning, and transmission electron microscopy. The parenchyma of the organ is predominantly red pulp, a system of splenic cords and sinuses. The white pulp consists of loose lymphoid tissue which forms a cuff around the pulp arteries. Filtering of particulate matter from the blood occurs in the red pulp by phagocytes of the pulp cords and ellipsoids (periarterial macrophage sheaths). The ellipsoids are pale-staining cuffs of macrophages and reticular cells in a framework of reticular fibres surrounding the arterial capillaries. Destruction of effete blood cells (especially erythrocytes) is confined to the pigment nodules; particulate matter is not taken up by the nodules. These yellow-brown bodies are dispersed throughout the red pulp and are bounded by a reticular capsule. They contain masses of phagocytes and have the appearance of a morula. They are associated with blood vessels and are surrounded by sinusoids. Prussian Blue stain shows the presence of haemosiderin within their phagocytes. The phagocytes of the pigment nodules are filled with inclusions such as residual bodies, siderosomes, and fragments of erythrocytes. The early filtering of particulate matter by the phagocytes of the pulp cords and ellipsoids may allow for a more efficient phagocytosis of erythrocytes by the pigment nodules, followed by storage and reutilization of iron-containing compounds uncontaminated by other phagocytosed material.     

Ultrastructure of the red pulp in spleen innervation in horse and pig.

The innervation of the red pulp in the spleen of horse and pig was investigated by electron microscopy. In addition, the neurilemma was studied by immunohistochemistry specific for S-100 protein. In the pig, a large population of smooth-muscle cells extending from the smooth-muscle trabeculae was present in the red pulp. The cytoplasmic processes of reticular cells enwrapped the smooth-muscle cells, and nerve fibres were distributed between the smooth-muscle cells and the reticular cells. The nerve terminals clustered toward the facing of the smooth-muscle cells. Nerve fibres and terminals were not disclosed within the sheathed artery. Immunohistochemically the neurilemma showed a reaction positive for S-100 protein. In the horse, no smooth-muscle cells were noted in the red pulp. The nerve fibres terminated around the cytoplasmic processes of the reticular cells. Nerve fibres and terminals were disclosed within the sheathed artery, and the terminals contained both large and small dense-cored vesicles. Immunohistochemically the neurilemma showed a reaction negative for S-100 protein. These findings support the presence of the axon-bearing reticular cells described earlier in the horse spleen.     

α- and β-receptor control of catecholamine secretion from isolated adrenal medulla cells

Summary The structural characteristics and cellular elements of the boundary zone between the white and red pulp of the human spleen were studied by SEM and TEM. The boundary zone consisted of both the perifollicular region and the region surrounding the periarterial lymphoid sheath. The perifollicular region was further subdivided into two, equally thick layers. The inner half layer of the perifollicular region outside the mantle zone of the lymph follicle was composed of tightly packed medium-sized lymphocytes, interspersed by a small number of reticular cells. The outer half layer was composed of a reticular cell meshwork containing blood cells in vessels, which communicated with the splenic cords of the red pulp. Intermittent rows of reticular cells distinguished the outer from the inner half layer. The region surrounding the periarterial lymphoid sheath revealed the same type of reticular cell meshwork as the outer half layer of the perifollicular region. Capillary ends opened into the reticular cell meshwork, which suggested the presence of an open circulation in the human spleen. A deep lymphatic vessel which communicated with the periarterial lymphoid sheath was noted.     

Microscopy of vascular architecture and arteriovenous communications in the spleen of two odontocetes

The red pulp of the spleens of the short-finned pilot whale (Globicephala macrorhynchus) and the Pacific bottle-nosed dolphin (Tursiops truncatus gilli) (Odontoceti) were examined by light and electron microscopy and found to comprise two venous layers, an inner and an outer. The inner layer is homologous to the intermediate zone (IZ) of primitive-type mammalian spleens and contains sinusoids consisting of endothelial cells and a thin layer of extracellular deposits. Its vascular structure is unclear. The venous vessels of this layer eventually communicate with veins of the perivenous outer layer. The perivenous layer contains veins of various sizes, interstitial elements, and trabeculae. It is filled with blood cells, particularly plasma cells, but no myeloid cells. The perivenous layer (PVL) is homologous to the red pulp of common mammalian spleens but shows signs of involution. The white pulp gives origin to arterial terminals that end in the red pulp, where they communicate directly with the sinusoidal veins producing a closed circulation. The arterial terminals do not show ellipsoids. The presence of an IZ with a closed circulation and the involution of the red pulp makes the spleen of Odeontoceti another example of a mammalian spleen of the primitive type that has been altered by the evolutionary process. Vascular remodelling of the spleen of Odontoceti seems to follow the pattern noted in the spleens of nonmammalian vertebrates. © 1994 Wiley-Liss, Inc.     

Histological, histochemical, and fine structural observations on the spleen of seals

Spleens of three species of Antarctic seals with different diving habits (Weddell seal, crabeater seal, and fur seal) have been studied with histological, histochemical, and electron microscopic methods. The spleens can be classified as nonsinusoidal, with capsule and trabeculae rich in innervated smooth muscle cells. The trabecular system is particularly well developed in the deep- and long-diving Weddell seal. As in other mammals the pulp can be divided into white and red pulp. In the white pulp, periarteriolar lymphatic sheaths and secondary lymphatic nodules occur; both are surrounded by a marginal zone rich in macrophages and eosinophils. The nodules can be observed frequently, which is in accordance with abundance of plasma cells in the red pulp. Well-developed white pulp and numerous plasma cells and eosinophils obviously reflect a high load of nematodes, which have mainly been found in lung and stomach. Additionally, in the red pulp morphological evidence for the following functions has been found: destruction of erythrocytes, erythropoiesis, and thrombopoiesis. In respect to blood flow through the red pulp, we interpret our observations in the following way: terminal branches of arterioles open into the space between the fibroblastic reticulum cells; blood draining from here is collected into pulp veins, which are mainly found near the trabeculae. Thus, the seals have an open vascular compartment in their spleens, as also occurs in the cat. The red pulp is innervated by numerous nerve fibers that seem to include both cholinergic and adrenergic ones. The target cells of these fibers seem to be the fibroblastic reticulum cells, whose state of contraction may influence the direction of blood flow through the red pulp.     

Ultrastructure of human spleen in transmission and scanning electron microscope

Despite new information concerning functional morphology of spleen, there are still some inaccuracies mostly regarding the spleen blood circulation. Billroth’s (splenic) cords are formed from three-dimensional network of fibroblastic reticular cells located among branched sinuses. Results from our study using scanning electron microscopy confirm an intimate contact between adjacent reticular cells and erythrocytes. Arterial terminals can be observed in the Billroth’s cords. The wall of sinuses reminds a sieve and it is lined with a special type of endothelium. In electron microscope, endothelial cells look like rods oriented parallel to the longitudinal axis of sinuses. Based on our observations fibroblastic reticular cells change to fixed phagocytes under no circumstances, hence they do not participate in phagocytosis. They may have a recognition function for cells circulating around them. According to our opinion, the open and the closed blood circulation are present in the human spleen simultaneously. Blood flowing in the closed circulation can help “absorption” of extra-vascular liquid and the blood elements into the vascular lumen. Due to sporadic occurrence of smooth muscle cells in the capsule and trabeculae, we assume that human spleen is not a blood reservoir, unlike the spleen in some other animals.     

The spleen and skin wound healing in Xenopus adults

In most vertebrates, the regenerative capacity to restore lost/damage tissues to original structure and functionality decreases at some time during ontogenesis. To evaluate the role of the acquired immunity in the decline of regenerative potential, we examined the cellular responses elicited in the spleen during skin repair in Xenopus adults. Modifications in the architecture were found to be induced and were remarkable 14 days postinjury when the spleen increased significantly in size. In white pulp, the periarteriolar lymphoid sheaths were associated with follicles having central light zones, morphologically similar to germinal centers. With the progress of healing, pigment‐containing cells were seen to accumulate in both white and red pulp regions. Moreover, compared to controls, the cells immunoreactive to anti‐cytokines (TNF‐α, TGF‐β1) and ‐iNOS increased from the first days after wounding. The 14th day, the positive cells formed a dense network of reticular cells in central regions of lymphoid follicles and more frequent reactive leukocytes were detected within the red pulp. A higher number of lymphoid cells immunostained with anti‐CD3ε were also observed in the perifollicular zone. The results suggest that the spleen of adult frogs is involved in skin wound healing with the expansion of lymphoid compartments. J. Morphol. 277:888–895, 2016. © 2016 Wiley Periodicals, Inc.     

The equine spleen: an electron microscopic analysis

The capacity of the equine spleen to store and rapidly release as much as half the circulating blood volume after adrenergic stimulation depends upon the size of the spleen, its muscular capsule, and the distinctive structure of its red pulp. The unit, or lobule, of red pulp is a cylinder of pulp spaces organized in a reticular meshwork, supplied by a peripheral ring of arterial capillaries, and drained by a central venule. Reticular cells, which make up the meshwork of the pulp, contain an extraordinarily large complement of microfilaments and intermediate filaments and are richly innervated by nerves containing both dense and lucent core vesicles typical of adrenergic nerves. The wall of the pulp venule contains large apertures. The capacious red pulp would thus appear capable both of large-scale blood storage and, by the contractile adrenergic innervated reticulum and open venous vasculature, of rapid expression of stored blood into the circulation. Antigen-presenting cells are present not only in B and T cell zones in white pulp but in the periarterial macrophage sheath of red pulp as well. The periarterial macrophage sheath is one of the first sites of antigen capture, and the presence of these cells confers on it an immunological role.     

Presence and release of the chromogranin B-derived secretolytin-like peptide KR-11 from the porcine spleen

Chromogranin B (CgB) is a major matrix protein in secretory granules/large dense-cored vesicles and a precursor for smaller peptides. In an earlier study, we have identified a secretolytin-like peptide (KR-11, pCgB(637-647)) from porcine chromaffin granules. Further evidence is presented here to show the processing of chromogranin B to this peptide during axonal transport in the splenic nerve and its release in the spleen upon various conditions of stimulation. Immunohistochemical staining showed that in the porcine spleen chromogranin B and NPY completely colocalize in nerve fibres and varicosities in blood vessel walls and trabeculae, and along the loose network of smooth muscle cells in the red pulp, as identified by their alpha-smooth muscle cell actin content. No antibacterial activity was found for the porcine secretolytin-like peptide, KR-11. The change of one amino acid residue (Thr-->Asn) in the porcine homologous fragment of secretolytin appears to be responsible for its loss of antibacterial activity.     

The marginal zone and marginal sinus in the spleen of the gerbil. A light and electron microscopy study

The spleen of the adult gerbil (M. unguiculatus) is characterized by the absence of venous sinuses and ellipsoid sheaths. The follicle (white pulp) is separated from the surrounding red pulp by a distinct marginal zone. The cell types in the marginal zone are common to both the follicle and red pulp. Separating the marginal zone from the follicle is a vascular channel of capillary dimension, the marginal sinus. A number of terminal segments of the arterial vessels within the follicle were observed to form a direct connection with the marginal sinus. Ultrastructurally, discontinuities were evident within the walls of the marginal sinus that would permit passage of both cellular and plasma components from the marginal sinus to either marginal zone or the follicle.     

Spleen mass as a measure of immune strength in mammals

• 1 In studies of birds and their pathogens, spleen size has frequently been used to make inferences about immune system strength. However, the use of spleen size in mammals is more complicated because, in addition to having an immune function, the mammalian spleen is also a reservoir for red blood cells.
• 2 To assess the reliability of mammalian spleen mass as an indicator of immune activity, we quantified the white and red pulp mass by histology of spleens from shot red deer Cervus elaphus. We then analysed the relationships among spleen mass, the amounts of white and red pulp, and the deer's body condition relative to faecal counts of the nematode parasite Elaphostrongylus cervi.
• 3 White and red pulp mass were positively correlated so that an increase in spleen mass was a positive function of both components of the spleen. In male deer, which had significantly lower body condition and higher parasite loads than females, parasite counts were negatively correlated with spleen mass, white pulp mass, and red pulp mass.
• 4 Our findings suggest that (i) spleen mass in shot red deer is a reliable measure of white and red pulp content; and (ii) when looking at the red deer life history, which is greatly influenced by sex of the deer, splenic mass and white pulp mass could be used as reflections of immune system strength.
• 5 Future studies of mammalian spleens can contribute to the understanding of evolved strategies of immune response investment in mammals. However, determination of the white and red pulp spleen components using various sampling methods must be made prior to their application.