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The Three-Dimensional Microstructure of the Liver a Review by Scanning Electron Microscopy

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The Three-Dimensional Microstructure of the Liver a Review by Scanning Electron Microscopy Scanning Electron Microscopy Volume 1986 Number 3 Part III Article 21 7-22-1986 The Three-Dimensional Microstructure of the Liver A Review by Scanning Electron Microscopy G. Macchiarelli University of Rome P. M. Motta University of Rome Follow this and additional works at: https://digitalcommons.usu.edu/electron Part of the Life Sciences Commons Recommended Citation Macchiarelli, G. and Motta, P. M. (1986) "The Three-Dimensional Microstructure of the Liver A Review by Scanning Electron Microscopy," Scanning Electron Microscopy: Vol. 1986 : No. 3 , Article 21. Available at: https://digitalcommons.usu.edu/electron/vol1986/iss3/21 This Article is brought to you for free and open access by the Western Dairy Center at DigitalCommons@USU. It has been accepted for inclusion in Scanning Electron Microscopy by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. SCANNING ELECTRON MICROSCOPY /1986/111 (Pages 1019-1038) 0586-5581/86$1.00+05 SEM Inc., AMF O'Hare (Chicago), IL 60666-0507 USA THETHREE-DIMENSIONAL MICROSTRUCTURE OF THE LIVER A Review by Scanning Electron Microscopy G. Macchiarelli and P.M. Motta* Department of Anatomy, University of Rome Italy (Received for publication March 13, 1986, and in revised form July 22, 1986) Abstract Introduction The improvement in scanning electron micros­ The structure of the liver reflects its ex­ copy (SEM) techniques has permitted us to des­ tremely complex function. In fact, the liver can cribe the microstructure of the liver. By SEM,the be considered as both exocrine and, to some ex­ liver peritoneal surface is composed of flat me­ tent, endocrine in nature (81). In addition, it sothelial cells possessing microvilli and cilia. metabolizes, detoxifies and inactivates many dif­ Hepatic sinusoids connect the portal vessels with ferent substances of both endogenous and exo­ the terminal branches of the hepatic vein (cen­ genous origin ( 46). It further plays an i mpor­ tral veins).Endothelial cells of the portal space tant role in the hemodynamic regulation of the arteries are elongated and arranged longitudinal­ splanchnic system. ly, while those of the central and portal veins When the first microscopists began studying are polygonal and flattened,possessing microvil­ the histological organization of the liver, it li. The sinusoidal endothelial cells show both immediately appeared clear that a visualization small fenestrations (sieve plates),up to 200 nm of its three-dimensional structure was funda­ in diameter, and large ones,up to l ~m. Within mental to unravelling its complex functional mys­ the sinusoids are seen bridging structures,cover­ teries. Several ingenious attempts to draw the ed by fenestrated endothelium,seeming to have a liver's three-dimensional pattern have been made fibrillar core. Kupffer cells resemble macropha­ (7, 15-17). Only in the second half of this cen­ ges,showing microvilli, blebs, lamellipodia and tury, however, through careful studies using ste­ filopodia. Within the Space of Disse are seen the reological methods and statistical analysis of fat-storing cells,having laminar dendritic pro­ transmission electron microscopic (TEM) pictures jections. The polyhedral liver cell faces the (18, 126, 145), has it been possible to confront Space of Disse (vascular pole) or faces an adja­ the heterogeneous complexity of the hepatic cent hepatocyte (biliary pole). Vascular facets gland. The limits of these techniques have been are evenly covered by microvilli. Biliary facets recently reviewed (73) and, at the present time, show a central longitudinal depression,bordered scanning electron microscopy (SEM) seems to be by microvilli (bile hemicanaliculi). Canaliculo­ the technique of choice for three-dimensional ductular junction and bile duct epithelia show studies. Thus, after more than 15 years since the blebs, microvilli and cilia. Up to now,fetal li­ first SEM reports of mammalian liver lobule (4, ver and liver pathology have been scarcely inves­ 23), so many SEMliver images have been obtained, tigated by SEM: in the future, they can be suc­ that we feel confident in our description of the cessfully approached by three-dimensional studies. "microstructure of the liver" (73). In fact, SEM has been extensively used to obtain data from the KEY WORDS:Liver, Sinusoid, Sinusoidal Cells, livers of many mammalian species including hu­ Hepatocyte, Biliary System, Pathology, Fetal, mans, and other animals, such as birds and fish Scanning Electron Microscopy. (see Table 1). Most of this literature will be reviewed in this paper. Further, new SEMimages of sinusoids *Address for correspondence: from an unexplored rodent (Praomys Mastomys Nata­ P. M. Motta, Department of Anatomy, University lensis) will be presented. of Rome, via A. Borelli 50, 00161 Rome, Italy. Phone No. (39-6) 49.26.23. 1019 G. Macchiarelli and P.M. Motta such as endothelial fenestrae (156, 157). Table 1. Primary fixation is usually performed with solutions of 1-3% glutaraldehyde in cacodylate or SOURCEOF LIVERTISSUE FOR SEMINVESTIGATIONS. phosphate buffer. Paraformaldehyde has also been (Bibliography update 1985) used, alone (4% in phosphate buffer), or with glutaraldehyde (1 .5 - 4% in phosphate buffer). No side effects have been demonstrated when the al­ Source Bibliography dehyde phase of fixation has been prolonged. This suggests that the samples may also be left in the CAT 62; 69-72; 81. primary fixative for a long period in order to CHICK 113. obtain optimal tissue fixation and hardening (5). DOG 69; 70. In a blood-rich organ such as liver, it is FISH 97. necessary to accurately wash out all blood from FROG 111. the vascular bed prior to fixation (81,116). In HUMANS 27; 36-40; 44; 62; 76; 81; 89; 91; this way, precipitation of plasma proteins which 134; 136-138; 153; 160; 161. obscure vessel surfaces, can be avoided (27). PIG 62; 70; 71. Several washing solutions have been used but no PRIMATES data can be referred as to which is the solution Baboon 57. of choice. Monkey 87; 108; 133; 135; 139. Both the addition of anticoagulants (hepa­ RODENTS rin) or vasodilatants (procaine), to infusates, Guinea-Pig 35; 62; 69-71. and the injection of either during anesthesia im­ Hamster 59; 160. proves the quality of perfusion, which can be Mouse 27; 69-71; 92; 121; 130; 141. monitored by noting the change in liver colour Rabbit 27; 105; 111; 125; 158. from a dark-red to a sandy-beige (116). Rat 8; 9; 11; 14; 21; 22; 26; 27; 30; Some authors suggest the maintenance of in­ 34; 41; 43; 45; 51; 54; 55; 62; 65; fusates at body temperature in order to mimic the 67-72; 74-79; 81; 86; 88; 90; 93; physiological environment and to enhance the 94; 99; 101; 112; 114; 115; 123; speed of fixative penetration (5). On the other 130-132; 142; 144; 149; 151-154; hand, the use of lower temperatures may reduce 158; 160; 162. anoxic damage. SHEEP 157. Attention must be paid to the infusion pres­ FETAL sure (153). In fact, the use of vasodilatant animals 71; 72; 76; 113; 157. drugs and aldehyde fixation through the arterial humans 58; 71; 76. circulation, has the effect of converting the blood vessels into a rigid pipe system, causing loss of the arteriolar "barrage" (22). Thus, the Preparation of Liver Samples for SEM infusates may reach the postarteriolar vessels (sinusoids) with a pressure higher than physiolo­ The improvement in electron microscopic gical values, which range from 6 to 10 mmHgin techniques during the past 15 years has permitted the mammalian portal system. In addition, there us to obtain optimal liver specimens for SEM is evidence that a high infusion pressure of in­ investigation with relatively simple procedures. fusates, whether injected via the portal vein or In order to avoid artifacts and to yield a clean via the arterial system, is associated with an surface for observation, however, it seems increase in the number of larger gaps in the si­ imperative to respect certain crucial points dur­ nusoidal endothelium (21). Considering certain ing the preparation of samples (27, 81). properties of endothelium, such as filtering and In order to obtain good preservation of hep­ sieving, these larger gaps have been also des­ atic subcellular structures, fixation must be cribed as artifacts (155) or pathological featu­ performed by vascular perfusion (148). In fact, res ( 21). immersion primary fixation alone does not ade­ In animal experimentation, perfusion pre­ quately avoid cellular anoxic damage, does not ferably is performed via the portal vein, but may preserve endothelium (150) and does not display a also be done via the aorta, or directly via the homogeneous fine structure in the whole sample left ventricle. Biopsy samples also have been (119).Furthermore, immersion fixation may create perfused, through the introduction of a small spatial artifacts due to the non-homogeneous needle into the parenchyma (puncture-perfusion) hardening of the liver tissue and it may also (38,40,86,134,137) or through cannulation of the cause the disappearance of delicate structures cut vessels (58,76,81). 1020 SEMof the Liver Osmication is not always required for SEM. useful to obtain stereo pictures during SEM In fact, it doesn't seem to offer any advantage observation, to allow a better understanding of during observation of liver samples (133). How­ the three-dimensional relationships of liver ever, the metal-impregnation technique with tan­ cells (73). nin-osmium may enhance the conductivity under the electron beam of non-coated specimens (84,85). The Peritoneal Liver Surface Dehydration can be carried out gradually, either with ethyl alcohol or acetone (81), fol­ The external surface of the liver is almost lowed by critical-point drying (see Howard and completely covered by a monolayer of serosal Postek, 1979 for further references) (28). cells that rest with the basal lamina on a vari­ Freeze-drying also has been performed, and is ad­ able amount of connective-tissue fibres, forming vantageous in exposing more of the surface area the fibrous capsule (73).
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