Influence Of Vagotomy At The Perfusion Of The Mucous Membrane Of The Duodenum And Jejunum With Blood
I. INTRODUCTION
The histological structures of the mucous membrane of the small intestine.
The micro-circulation of blood of the mucous membrane of the small intestine.
The lymphatic circulations of mucous membrane of the small intestine.
The sympathetic innervation of the intestinal uvulas.
The sympathetic stimulation and micro-circulations of the mucous membrane of the small intestine.
The model of the circulation of the mucous membrane of the small intestine in rabbits.
Calculation of the flow of blood toward segments of circulation.
The absorption’s model of the mucous circulation of the small intestine.
The clinical importance of the dissertation.
II. MATERIALS AND METHODS
Experimental animals.
Operational process.
Calculation of flow.
III. RESULTS
Variability of the mucous flow. Probability.
Comment of results.
Initial post-vagotomy's reaction of the duodenal mucous
membrane and jejunums.
Influence of a hypoxia on the post-vagotomy's circulation
of the duodenal and the jejunal mucouse membranes, and
vice versa.
Influence of a hypercapnia on the postvagotomical
reaction.
IV. DISCUSSION
V. CONCLUSION AND SUMMARY
VII. LITERATURE
VIII. SUPPLEMENTS
The goal of this work is an answer to the question, whether, and under which conditions, a vagotomy can stop bleeding out of the mucous membranes of the duodenum and the small intestine. It is known that vagotomy opens a-v anastomosis in the mucosa and submucosa of the stomach producing an ischemia of the most superficial layers (1, 2,). Spanner described the existence of such anastomosis also in intestinal mucosa of men, rats and in submucosa of dogs (3). Bell and Battersby with clearance of radioactive Krypton-85 proved that vagotomy reduces the blood flow through the mucous membrane of stomach for 29 -74 %, depending upon completeness of intervention and reactions experimental animals (4). So they confirmed works of others authors that vagotomy reduces the total blood flow (5, 6,) and the mucosa blood’s volume (2). Vagotomy prevents grow the blood flow after stimulation of the front hypothalamus (7). On the base of these and others understandings were made the first both-sides truncal vagotomies with piloroplastics for stopping massive bleeding from the most superficial layers of gastric mucous membrane (8).
In the master's thesis: “The modified selective vagotomy as a supplement to surgical treatment of erosive gastritis" I presented my experiences with implementation of vagotomy in the treatment of massive bleedings caused by acute erosive ulcerations of the gastric mucous membrane. (Zagreb, 1981.)
From the large scientist interest is the influence of vagotomy on the perfusion of duodenal and jejunal mucous membranes. Therefore, we decided for an experimental scientific investigational work from this area. How frequently, concomitantly in the same time together with the gastric mucous membrane, bleeds also the duodenum and the jejunum (erosive gastroduodenitis and gastroduodenojejunitis), investigation of influence of vagotomy on the perfusion of mucous membrane latter has alsop the clinical importance. If it is identical that on the mucous membrane of stomach "The modified selective vagotomy” could be an operation for the bleeding from intestine too.
I. INTRODUCTION
For the understanding of this work is necessarily to get acquainted with the histological structure of the mucous membrane of small intestine, her microcirculation and the sympathetic innervations, especially with the influence of sympathetic stimulation on the redistribution of blood flow and models of circulation at rest, ingestion, digestion, strengthened motility and other. With comparison of the known with results of our investigations, we will carry out conclusions about the influence of vagotomy on the perfusion of the duodenal and the jejunal mucous membrane.
II. HISTOLOGICAL STRUCTURES OF THE MUCOUS MEMBRANE OF SMALL INTESTINE
The wall of the small intestine makes mucous membrane, submucosa, muscular layer and serosa. The last three layers are permeated with unique system of crossing collagen fibers.
THE MUCOUS MEMBRANE consists of epithel, lamina propria and muscularis mucosae.
CIRCLE’ WRINKLES (plicae circulares Kerkringi) are best developed in the distal part of the duodenum and proximal the jejunum, while in the ileum they become gradually all smaller and thinner, and gradually disappear. They are built of the mucous membrane and one part of the submucosa.
INTESTINAL UVULAS (villi intestinales) cover complete internal surface of the small intestine. They are long 0,5 to l,5 cm, dick 0,l mm. They are outgrowth of epithelium and laminas propria. They freely stick out in the lumen. On 1 mm² of surfaces have them around 30, total several millions. Between uvulas are opens of intestinal glands.
EPITHELIUM of the small intestine is single-layer with two type cells. Most numerous are cylindrical or resorption’s cells. Less numerous are jug’s cells. Cylindrical cells are high about 20 micrometers (microns). They are very elastic, and can easily adapt to all changes. Theirs cytoplasm is finely granular or honeycomb, towards theirs functional’ condition. Their core has placed in the basal part of cells. Theirs surface is covered with 1 -1,5 mm of the fat coating that consists densely compressed shoots of cytoplasm (mikrovili). They are long 1 mm, with the diameter about 0,1 µm (the micrometer). They are increasing the surface of the cylindrical (resorption’s) cells about 14th times. In every mikrovila enters a zone of microfilaments, which occupies theirs centers.
The jugs’ cells are put among cylindrical cells. In the jejunum are they less, in the ileum more. On the free surface, they have no coatings.
THE epithelium with intestinal uvulas enters in Lieberkühn’s crypts, i.e. intestinal glands (glandulae intestinales) and coats them. Those are simple tubular glands long 0-2 to 0,5 mm, placed in the lamina propria. Cylindrical cells are in them are something shorter of those on the surface of intestinal uvulas and they are constantly mitotic dividing, enabling the regeneration of the intestinal epithelia.
For Lieberkühn’s crypts in the small intestine are characteristic Paneth’s cells, which are coming individually, or in groups, on theirs bottoms. They have a round core. Theirs cytoplasm are filled with numerous acidophil granules. Ultra-structurally, at the surface, can be seen mikrovili. At the bottom of Lieberkün’s crypts, there are still yellow cells, which characterize fine granules in the basal part of cytoplasm, which stain with chromic salts yellow. Yellow, enterocromaffin, or basal granulated cells, are belonging to the group of endocrine cells of the gastrointestinal system. In different parts of the digestive system they are intended to various secretor roles (they secretes serotonin, gastrin, secretin, pankreozimin and enteroglukagon). Some are placed in the base of the intestinal epithelia and have no link with the lumen (endocrine cells of the closed type), and others are connected wit the lumen (endocrine cells of the open type). These last possess mikrovili, which can be very long and which receive informations from the lumen on which basis they secrete theirs granule in surrounding capillary spaces. Some of those endocrine cells are morphologically and functionally related to cells of Langerhans’ little islands of pancreas, so a part of researchers count these cell in the unique “gastro-enteral-pancreatic” (GEP) system.
THE LAMINA PROPRIA of the small intestine fills uvulas and the space between Lieberkühn’s crypts. It is built of the reticulated tissue, which contains many free cells. On some places, it is filled with a dense mass of lymphocytes, so we speak about the lymph tissue, which comes in two forms: as single lymph little knots (Payers’ plates in the ileum). Besides it, lamina propria contains large number of plasmas’ cells, eosinophilic leukocytes, an some mast-cells. Special kind cells of the laminana propria are so-called globular’ leukocytes, which are emphasizing with theirs round granules, or beads, which color with eosin red. Men give them immunity’s importance.
THE MUSCULARIS MUCOSAE enables moving the mucous membrane of small intestine. It comprises of two layers of the smooth muscular cells: interior’s mostly circular and outer longitudinal. Between bundles of smooth muscular cells is a network of fine elastic fibers, which crosses in the propria. Smaller bundles of the smooth muscular cells enter in the propria of intestinal uvulas and inside it lay down parallel with the longitudinal axle. With its contraction, the uvula shortens and pushes the blood and the lymph from them in larger blood and lymph vessels.
THE DUODENUM has very tall circle wrinkles with dense thickset uvulas on the surface and special kind of glands, which there are not in others parts of the small intestine. That are Brunner glands – the duodenal glands (gl. duodena1es). They are built like the gyrate ramified tubular glands. With larger part they go through muskularis mucosa and fill out the larger part of submucosa. Their drainage canals open in intestinal crypts, or between uvulas. Brunner glands are built of mucous cells similarly to pyloric glands. They secrete bright mucous excrete which contains proteolytic ferments, and participate at splitting of carbohydrates. Ultrastruktural contains a rich endoplasmic small net, numerous mitochondria, very large Golgy system, and numerous secreting granules of which ones are zymogene and the second mucous.
THE JEJUNUM. In the upper part of jejunum circle wrinkles are high and numerous, similarly in the duodenum, but towards the ileum they become shorter having all bigger interspaces. Uvulas are long and some more flattened. In the epithelium of the distal part of the jejunums jug cells are especially numerous. The lymph tissue is represented in the form of little single lymph knots in the mucous membrane (9).
THE VILLUS INTESTINALIS is the small protuberance like lobule. It is high 0,5 -1,5 mm. It is spread out widely whole the small intestine. It is built of the connecting basis, which makes the skeleton of uvula. In this basis have a lot of lymphocytes and some individual leukocytes. In the middle of the uvula is the central lakteal. Around it are two or more arteries (central arterioles), which go towards the top of the uvula, parallel with the central lakteal. On the top of the uvula arteries cross in a capillary network, and one twig goes directly in the vein as arteriovenous anastomosis. This anastomosis is important for functioning of the heart, because in no-digestive phase blood directly over the anastomosis goes in the vein and does not spend any energy for pushing blood through the capillary system. The smooth muscular fibers of uvula come from the muscular layer of the mucous membrane, go parallel with the longitudinal axle of uvula, and come to its top where they catch for the basal membrane of the epithelia. When this muscular fibers contract it empties and the again erects by the blood (9).
THE MICROCIRCULATIONS OF THE MUCOUS MEMBRANE OF THE SMALL INTESTINE
Large central arterioles of the villuses (intestinal uvulas) start from the submucosa’ arteries, and go parallel with the central lakteal (the hilus’ sinus) towards the top of the villus without branching. Nearby the top every the central arteriola gives two branches. The one as pre-capillary arteriola go parallel with the curve of the top and crosses in the subepithelial capillary net, which connects with the subepithelial capillary layer of marginal arterioles. The second, something distanced of the top of the uvula, crosses in the central vein. Capillaries of the upper branch go through the capillary subepithelial layer from the top of the uvula towards the base flowing into, in middle parts of the uvula in the central vena. Through this direct arteriovenous anastomosis between the subepithelial capillary layer and the central vena empty also capillaries of the subepithelial layer of basal parts of the villus which is continuation of the capillary layer of the region of crypts, in which blood flows from the base towards the direct arteriovenous communication in the middle part of uvula (2,111).
Two or more small branches of submucosal arteries penetrate the muskularis mucosa and enter in the region of crypts. Here, form the capillary net, which extends up to basal parts of the villus continue in the subepithelial capillary layer. Less pre-capillary arterioles go directly towards the subepithelial capillary layer as marginal arterioles (2, ll).
THE LYMPH CIRCULATIONS OF THE MUCOUS MEMBRANE OF THE SMALL INTESTINE
The absorbed liquid accumulates in the interstitial tissue where from blood and lymph capillaries take away it. Hydrostatic and the oncotic pressure have the crucial role in the drainage of this liquid. The drainage ability of these systems are fascinating (13).
The model of blood and lymph microcirculations, shown in the Fig. 7, shows some details that characterized the architecture of the villus.
1. Villus’ capillaries are 0,5 – 2,0 micrometer of the basal membrane of the epithelial cover. They create the capillary subepithelial layer thick 2,0 µm.
2. The fenestrations of the capillary endothelia are on the side of the basal membranes and communicate with the subepithelial and the juxtacapillary interstitial lymph space.
3. The central lakteal is located average 50 micrometers of the basal membrane of mucous epithelial layer.
4. The space within the villus, between arteries, represents the interstitial lymph space in which collects absorbed liquid.
Many think that via subepithelial juxtacapillary space takes away majority of absorbed liquid. (The lymph capillary layer is analogous to the blood subepitelial capillary layer.)
Accumulation of liquid in the interstitial tissue leads to:
1. Growth of the volume of the interstitial liquid;
2. Growth of the interstitial hydrostatic pressure, and
3. to decrease of interstitial of the oncotic pressure.
The interstitial tissue is composed of the collagen threads and mucopolysacharides mutually interwoven in the gelatinous mass. The hydraulic conductivity normally hydrated interstitial tissue is very small. As the volume conductivity vice versa proportional to concentration of the hyaluronic acid, the growth of the volume of liquid increases the hydraulic conductivity of the interstitial tissue. The hydrostatic pressure increases too. The oncotic pressure falls. The capillary filtration stops. Filterable capillaries turn into absorptive. The lymph’ vessels are filling with the liquid. During absorption oncotic lymph pressure falls with 1,3 kPa (10 mm Hg) – normal – on 0,3 to 0,9 kPa (2 to 7 mm Hg). Fall of the interstitial oncotic pressure for 0,3 to 0,9 kPa (2 to 7 mm Hg) depends on the level of water absorption. Conductivity of the interstitial tissue is, therefore, larger what its volume is bigger. (The oncotic pressure is reduced). By it the capillary conductivity and permeabilitet remains unchanged.
Divisions of the interstitial tissue on central and the juxtacapillary part ensure the passive flow of liquid in capillaries. The hydrostatic pressure is in juxtacapillary areas much more sensitive on a change of the volume in the central interstitial spaces of the villus. Therefore, larger part of the absorbed liquid takes away hereby and this passively in capillaries. The largest fall of the interstitial oncotic pressure appears just in that subepithelial-juxtacapillary part.
40 % of absorbed liquids are taking away with lymph vessels, dependently on:
1. The pressure of the intra-lumen liquids in lymph vessels;
2. The portal pressure;
3. The mobility of the small intestine and
4. The blood pressure of arterioles and capillaries.
The interstitial hydrostatic pressure moves liquid through the tissue and large pores of walls of limph vessels. The flow of lymph maintains valves of lymph vessels and the gradient of the hydrostatic pressure. Of help is also active contractions of arteries (inside factors) and the mobility of intestine, and contraction of villus (outer factors). The fall of the pressure in the central lakteal and collection of accumulated vessels makes easier filling (the distension pressure). Gastro-intestinal hormones (cholecystokinin and secretin) instilled in the a. mesenteric superior are increasing the intestinal flow of bloods and lymph. Turner and Barrowman think that hormones exuded during absorption increase the capillary perfusion and the pressure in capillaries. Increased capillary filtration increases amount of liquid in the interstitial tissue. Cholecystokinin and secretin would so accelerate the flow of lymph, preventing transforming exuding capillaries in absorptions’.
The amount of the absorbed waters via lymph roads can vary from 1 to 85 %.
At the normal portal pressure, blood capillaries remove the larger part of absorbed volume. Blood capillaries fills also the hydrostatic and oncotic pressure, while lymph only hydrostatic. At increased the portal pressure is reduced evacuations via blood. The hydrostatic pressure grows, as also evacuation with lymph. Because of a stoppage are growing capillary filtration. Lymph vessels remove also the additional interstitial volume.
After meals or intralumenal applications of nutrients increase the interstitial flow for 30 up to 130 %, causing a postprandial intestinal hyperemia. The absorbed liquid increases the capillary hydrostatic pressure. This, on the other hand, reduces the pressure of the interstitial tissue on capillaries. The oncotic pressure of plasma falls. The capillary’s perfusion and permeabilitet grow. In the absorptive phase, therefore, absorbed liquid spreads the interstitial space, increasing the hydrostatic pressure and reducing the oncotic pressure. Changes are causing a reverse flow: from interstitial tissue in the blood. Filterable capillaries convert in absorptive. Grows production of lymph and accelerates the lymph flow (13, 14,).
SYMPATHETIC INNERVATIONS OF THE INTESTINAL UVULA
Noradrenergic fibers flow along blood vessels. The muskularis mucosa and the submucosa are especially well innervated. Dense plexuses surround arterioles in the region of crypts from which threads ascend in the hillus. Together with central arterioles, they go along the central lakteal. On the top of the uvulas, they branch, creating dense tangles of subepithelial capillary layer of the villus top and base-lateral membranes, whose terminations terminate in capillaries. Noradrenergic threads are found also between epithelial base-lateral membrane and capillaries. However, these fibers never noticed beside the epithelium. Exceptionally they could be viewed together with capillaries (15).
Absorptions of liquid is under influence of noradrenalin. Alpha-adrenergic agonists (Field and Mc Coll, 1973), exogenously given noradrenalin (Fields and Mc Coll, 1973, Hubel 1976, Levens and coworkers 1979), and stimulations of the simpathetical threds in jejunum (Brunson and coworkers 1979) increase absorption of liquid. Alpha-adrenergic antagonists interrupt with noradrenalin-encouraged absorption (15).
Noradrenalin directly influences on the ion’s pump on the epithelial base-lateral membrane (Fields and coworkers 1976). It also influences on the flow of blood in the area of crypts, and in the villus. Probably it influences also on the lymphatic system of the villus with contraction of smooth musculature. Noradrenalin liberated in the tissue around capillaries diffuses in this, and via the blood comes on the second places, even in cells of the epithelium, which alone are not noradrenergic innervated (l5).
The smooth musculature of the villus is well noradrenergic innervated.
Levens and coworkers (1979) are opinion that alpha-receptors on base-lateral membrane of the epithelial cells
tie noradrenalin liberated at terminations and accelerate transport of liquid. (Results are identical those got after implementation of angiotensin II (Levens and coworkers 1981).
Blood vessels of the region of crypts are very well innervated, as well as connections capillaries and varicoses in vicinity of the base-lateral membrane. Threads, which accompany the central lakteal, have the crucial influence on the smooth musculature (15).
SYMPATHETIC STIMULATIONS AND MICROCIRCULATION OF THE MUCOUS MEMBRANE OF THE SMALL INTESTINE
Stimulation of the noradrenergic fibers of the villus
lowers lymph flow, coefficient of capillary filtration, and the capillary pressure. Also reduces the pressure of the interstitial circulation. This means that during stimulations of the sympathicus there is no "self-regulation" no one of these elements (l6, 21). Stimulations of fibers of the small intestine promptly reduces blood flow, which after gradually returns in normal frames. Arterioles and venulas contract balanced actively. Because of these biphasic answers come up to relative weak fall of capillary pressure. Intestinal flow promptly falls on 48,3 beside oscillations about 7 %. (Later increased on 73,7 with oscillations of 4,2 %.) Coefficient of the capillary filtration reduces for 75 %.
The adrenergic stimulation increases the total intestinal resistance for 47 %. Grows are caused wit increasing of the pre-capillary resistance. Growth of post-capillary resistance is insignificant. During sympathetic stimulations grows the lymphatic oncotic pressure, which indicates the intestinal dehydration (l7).
The sympathetic nerve system plays an important role in reflex control of the blood volume. The reflex activation of the sympathicus in the answer on hemorrhage causes vasoconstriction and grows pre- and post-capillary resistance. As, result of segmental changes of vascular resistance capillary pressure falls, and the interstitial liquid is absorbed in the vascular system for purpose of restoration of blood volume (the "auto-transfusion") (18).
Maintenance approximately constant capillary pressure during stimulations of the sympathicus considers important for protection of delicate function of the alimentary canal in the preventing intestinal dehydration (19, 20,).
Tonus of pre-capillary sphincters seems that plays
deciding role in reduction of capillary flow (l8). Reduced transport’s capacity of the intestinal microcirculation lows filtration.
Lundgren and coworkers have noticed growth of the absorption of liquid and electrolytes during stimulations of the sympathicus (18, 20, 22,). Others authors (Shepherd AP and coworkers) have confirmed that sympathetic stimulation closes pre-capillary sphincters. Decrease of capillary flow
lower giving oxygen to tissues on the level of diffusion (23, 24).
Some works (Gidda JS and coworkers) show that vagotomy has any effect on the electric activity of the small intestine. Conclusion: n. vagus influences excitatory and inhibitory. Inhibitory influences are dominant in an intact intestine (25).
MODEL OF THE MUCOUS CIRCULATION OF THE SMALL INTESTINE AT RABBITS
How our experimental work is worked on rabbits we must get acquainted with the model of theirs circulation. It is practically identical to man’s.
The theoretical model of vascular net the small intestine in rabbits has been built by injecting with the silicone foam and polystyrene micro-granules (Lewitt of DG and coworkers, 1979). The same authors have confirmed with this an earlier opinion that the blood flow in the area of crypts is independent about the flow in villuses. This was completely understandably because those are two areas with different functions (26).
Big central arterioles start from sub-mucous arteries and accompany the central lakteal to the top of the villus, or near it, without branching. Two or more branch of sub-mucous arteries go through muskularis mucosa and enter in the region of crypts. Here form the capillary network, which communicate with the sub-epithelial capillary net of villuses. This network belongs to marginal arterioles which branch in the capillary sub-epithelial layer.
Near the top of the villuses, or in alone tops, the central arterioles give two branches: one pre-capillary arteriole, which follows the curve of the villus branching in the capillary sub-epithelial layer of the top. Over the capillary networks, this layer connects with marginal arterioles –sub-epithelial capillary layer of the basal and the middle parts of the villus. The second branch of the central arteriole is something more remote of the top and crosses in the central vein. This is a -v connection on the top ("by-pass"). Capillaries of the first branch empty through direct a-v communications in the central vein in the middle part of the villus. Through these communications are emptying the capillary sub-epithelial layer of lower parts. This means that the circulation in the capillary sub-epithelial layer of upper parts of villuses go in the direction from tops toward the base, and in the distal parts from the bases towards tops.
(At the rat the central arterioles give two pre-capillary arterioles which crosses in the sub-endothelial capillary layer. There is no direct a-v communication on top.)
At the value of the pressure of 13,3 kPa (100 mm Hg) in the aorta, the pressure in arteries on relation aortas-intestine falls on 7,6 kPa (57 mm Hg). In the intestinal wall in intramural arteries measures 20 micrometers falls on 0,8 kPa (6 mm Hg), in order to then gradually increases as follows: on relation on mucous arteries measures 20 micrometers 2,0 kPa (15 mm Hg), in order to in mucous arterioles would have been 2,9 kPa (22 mm Hg).
CALCULATION OF THE FLOW OF THE BLOOD IN SEGMENTS OF THE CIRCULATION
Lewitt DG and coworkers are calculated the blood flow in segments of circulation. They used POISEUILLE law. Beside the constant viscosity the segmental resistance is directly proportional relationship to length, and reversely proportional to the radius on fourth.
l = the length
r = the radius
The segmental resistance = l/r on 4 (radius on fourth).
The segmental resistance is therefore directly proportional to length, and reversely to radius on fourth.
Length and radius of the blood vessels was determined microscopic. It was studied 50 villuses. (The Rabbit had the weight of 1 kg.) The average height of villuses was about 500 micrometers (400 – 800 micrometers). Radius of central vein was between 5 to 8 micrometers. Radius of central arterioles was about 5 micrometers, terminal arterioles and capillaries 2,5 to 4,0 micrometers.
Distributions of mucous flow in the model were calculated by analysis of the circulation’s networks under the microscope.
Throughput was determined from the value of the blood pressure and the segmental resistance in intramural veins measures 20 micrometers. The mucous pressure was 2,9326 kPa (22 mm Hg). The absolute mucosa throughput through villus was 5,37 x 10 on -5ml/min (ten on the minus 5). Number of villuses in a gram intestine was 2,2 x 10 on fourth. The mucous throughput through the gram intestine we can get if the absolute mucous throughput through one villus multiply with number of villuses in the gram intestinuma. It is 1,18 ml/min. The intestinal flow we can get if to this worth we add the non-mucous intestinal flow in the gram of tissue (5 % of the intestinal flow under normal conditions). It is 1,24 ml/min.
All is calculated on the normal arterial pressure of a rabbit of 12,7 kPa (95 mm Hg). The presumable pressure loss through the system circulation is 0,9 kPa (7 mm Hg). l2,7 kPa minus 0,9 kPa is 11,6 kPa. (95 mm Hg minus 7 mm Hg is 88 mm Hg.)
By the calculation of the segmental flow was used the relationship: Pressure = flow x resistance.
The throughput in the region of crypts is 44 %, and in the region of villuses 56 % of the total mucous flow.
With contraction of the central arteriole reduces its diameter. The capillary pressure falls. For example, by the decrease for 30 %, the capillary pressure in the theoretical model of circulation falls for 1,2 to 1,5 kPa (9 to 11 mm Hg). In the same time the mucous flow falls with 1,l8 ml/min on one gram of the intestine on 0,81 ml/min/gr. The throughput through the region of crypts grows with 44 % on 60 % of the mucous flow. The throughput through the region of villuses falls with 56 % on 38 % of the mucous throughput. Pressure in central and marginal arterioles grows with 3,9 kPa on 4,4kPa (with 29 on 34 mm Hg). At the mucous throughput of 0,81 ml/min/gr intestine, the intestinal throughput is 0,90 ml/min/gr. Difference refers on the non-mucous intestinal flow and on growth of the capillary pressure for 0,13 kPa (1 mm Hg) or less, what is additional time’s effect, because of changes of the blood pressure in the smallest arteries.
The absolute resistance of the mucous throughput calculated with Poseuille law, respecting known sizes, is 54653,0 kPa (4,1. x 10 on 5 mm Hg). Pre-mucous resistance is 40 % of the absolute: 21861,2 kPa (1,64 the x 10 on 5-in the mm Hg).
The resistance of branches of the central arterioles is 98642,0 kPa (7,4 the x 10 on 5-in the mm Hg), marginal arterioles 239940,0 kPa (1.8 the x 10 on 6 in the mm Hg).
For the mucous perfusion are importance resistances:
l. Central arterioles;
2. Marginal arterioles;
3. Capillaries;
4. Pre-capillaries arterioles and a-v connections;
5. Central veins.
56 % of the total mucous throughput enters mucosa via the central arteriole (through villus). Two-thirds of it (38 % the total throughput) go via "by-pass" on the top of the villus in the central vein. Only a third (1.8 % of the total throughput) runs through sub-epithelial capillary network. 44 % of the mucous throughput goes via marginal arterioles. The very small part relates on the circulation in the region of crypts.
The throughput in every sub-epithelial capillary area (4 distal steps of the model) is 2-4 % of the total flow. The largest part relates on upper steps – the top of the villus. The villous capillary throughput is uniform – equal in all villuses.
The throughput depends on the segmental resistance. The largest part of the total mucous resistance relates on the arterioles. If the resistance in central and marginal arterioles reduced on 0 (widely dilated), and the pressure stays the same, the throughput grows for 139 %. If the resistance falls in all else vessels except arterioles the throughput grows only for 80 %. If we reduce the resistance in the mucous throughput, the perfusion increases only for 33 %. In order to reduce the perfusion more than for two-thirds is necessary strong contractions all trio of arterioles, majorities capillaries (with open "by-pass"), with relating contraction of the central vein.
Three quarters of the total intestinal resistances relates to pre-mucous blood vessels. Therefore, the throughput of the mucosal resistance is not altering the perfusion as much as a growth of the pre-mucosal resistance. Shutting of the CA in the region of crypts reduces the mucosal throughput for 54 %. If the CA is closed in the region of villuses, decreasing is only 2l %. Closing of marginal arterioles in the region of crypts reduces the mucosal throughput for 43 %. Increasing or decreasing of resistance in the CA has the large effect on the villus throughput, and very small in the region of crypts. At the marginal arterioles are vice versa.
If the resistance in the CA reduced from 16 (normally 1) on 0,2 (the diameter grows of 0,5 to 1,5; normally 1), and others resistances remain unchanged, the villus capillary throughput grows rapidly from 37 % on 147 % (the normal flow 100 %). If the resistance in the marginal arteries reduced in the same proportion the growth of the villus capillary throughput is considerably smaller.
THE ABSORPTION MODEL OF THE MUCOUS CIRCULATION OF THE SMALL INTESTINE
Postprandial hyperemia is the complex phenomenon. It is caused with neural, humoral and local factors. In phases of the ingestion is noticed a prompt mesenteric vasoconstriction. During digestions, postprandial, the blood throughput gradually grows, in order to reach maximum about one hour after ingestion. It maintains for hours after. The blood throughput is doubled. Beside mucous arterioles are dilated also all else arteries (27).
Atropine weakens the postprandial hyperemia (27). A blockade of alphas- and betas- receptors does not disturb the mechanism of the postprandial hyperemia (27). Cholinergic blockade weakens vasodilatation. It also prohibits the excretion of intestinal hormones. Vagotomy prohibits excretion of these hormones. Directly after the operational intervention, vagotomy weakens postprandial hyperemia.
A fatty meal, in the experiment whole milk or the wheat oil, causes the hyperemia not just of windings in which the meal is instilled, than in neighboring windings too. The model of the circulation is similar to that, which we get after intra-arterial infusion of secretin or the cholecystokinin. The food, which contains a bit of greases and proteins cause a hyperemia only in the winding in which is applied (27).
With instillation of 5 % and 10 % the solution of glucose in 9 % the NaCl we get the model of the metabolic intestinal mucosal hyperemia. (Sodium enables an active transport of glucoses.)
The local anesthesia of the examined windings weakens the hyperemic effect intra-luminal applied glucoses. This suggests the nerve influence on the postprandial hyperemia. Hyperemic answer depends also about osmosis possibility of meals. What this is bigger more strength is hyperemia (27).
With motility induced hyperemia, as well as at the skeleton’s musculatures, causes the hyperemia of the muscular layer of the mucosa (27).
Absorbed liquid reduces concentration of intestinal proteins. Trans-capillary concentration’s gradient grows. Also accelerates circulation of lymph (27).
THE CLINICAL IMPORTANCE OF THE WORK
Hemorrhagic duodenitis and jejunitis are not rare how it is thinking. The mucous membrane of the small intestine often bleeds together with the mucosa of the stomach. Causes of the hemorrhagic lesions both are the same. Upper parts of gastrointestinal tract frequently bleed at chronic renal diseases (28). Some medicines can cause fatal bleedings: tolazolin for example in the neonatal period (29}. Cirrhotic patients also gladly bleed. Some racial groups are specially disposed toward the hemorrhagic enteritis. So is described the special clinical entity at Turks (30). Parasitic diseases are the further cause of hemorrhagic enteritis (31). Described is cases of the massive bleeding from the gastrointestinal tract at Recklinghausen disease (32). Oldsters generally bleed more easily than young people. The protein loosing enteropathy often finishes lethal because of the massive bleeding in mukoe the course of the erosive jejunoileitis (33). Very dangerous bleeding duodenitis and jejunitis are described after operational interventions on the spinal marrow and large blood vessels (34, 35, 36, 37, 38, 39, 40,). A massive intestinal bleeding can also cause yersinia pseudotuberculosis (41).
Stopping the massive bleeding from mucosal lesions of the small intestine is the big problem. Extensive resections are not permit. Therefore possible stopping such bleedings with vagotomy would be of the big clinical importance.
II. MATERIALS AND METHODS
Experimental animals.
After Lewit DG and coworkers, 1979 years, were built the theoretical model of vascular network of the mucosa of the small intestine of rabbits, it became an experimental animal of choice for studying the mucous circulations at men. CA gives, namely, as in men, two branches: one that makes the direct arteriovenous connection between the CA and CV, and the second that, as a pre-capillary arteriole crosses in the sub-epithelial layer. At the rat, CA gives two pre-capillary arterioles, which both crossing in the sub-epithelial capillary layer. Lewite DG and coworkers calculated also the throughput of the blood in segments of the circulation. By that, using Poiseuille law, on which, beside the constant viscosity SE (segmental resistance) = l/radius on 4 (l =length). The length and radius of the blood vessels they determined by microscope. They used a rabbit of 1,0 kg. They studied 50 villuses. The transversal height of villuses was 500 micrometers (400 - 800 µm). Radius of CV was between 5 to 8 µm, CA around 5 µm, terminal arterioles and capillaries 2,5 to 4,0 µm. Distribution of the circulation was presented in the introduction. .
This conceives enabled calculating the blood throughput in segments of the circulation in the same model and after vagotomy. Therefore, the rabbit is the experimental animal of this work. It wereused rabbits of 3,0 kg with the Department for the pharmacology, which otherwise were used for student’s exercises. Reactions of these animals on the pharmacy-drugs of the autonomous nervous system and on hormones of the gastrointestinal tract secretin and the cholecystokinin are identical as at man (14).
The work was made on 16 experimental animals. Ten experimental animals were used for the creation of main work, and others six for the testing of the initial reaction, and an influence of vagotomy on hypoxic and hypercapnic circulation. Tested was also the influence of vagotomy on the hypovolemic circulation, as on the redistribution of the throughput. Obtained results enabled studying of the influence of hypoxia and hypercapnia on the post-vagotomic circulation. Presented is the mucous throughput 1, 3, 10, 20, 30, 45, 60 and 90 minutes after vagotomy.
Operational procedure.
Animals were narcotized with 25 % solution of Urethan. It was given 0,9 mg on 1 kg of bodyweight intravenous in the vein of an ear. After introduction of an animal in narcosis followed a cut through the derma, subcutaneous tissue and pre-tracheal fascia, with what was presented trachea, through that was brought a permanent plastic cannula for a undisturbed breathing. The venter was opened with the median laparotomy. At all animals double-sided trunk’s vagotomy was performed. After mobilization of the abdominal oesophagus, this was completely cut several centimeters above the cardia and removed in the length of 1 to 2 cm together with the adjacent oesophageal tissue and both vaguses.
Samples of the duodenal and jejunal mucous membrane and were taken in certain intervals before and after vagotomy. Supply and drains’ blood vessels were occluded. In this way was prevented a discharge of the circulative volume by taking of samples for histological investigation. 15 - 30 minutes after putting samples in the fixative liquid (10 % solution of formalin)
clamps were removed, and the sample (in the whole circumference of the length of 2 - 3 cm) were cut on two half because better fixation. In the purpose of presenting of an influence of vagotomy on hypoxic and hypercapnic circulation of an isolated winding of intestine, and vice versa, was used a device for the extracorporeal circulation.
By help of the device for the extracorporeal circulation, in a canulated a. mesenterica cranialis, was applied the solution of colored gelatin the same viscosity with blood (500,0 ccm physiologic NaCl, 50,0 gr of gelatin and 30.0 gr of "intensive chine ink"). Used liquid was previously warmed on 37°C.
The histological preparations were made in the Department for the histology of the Medicine faculty in Zagreb and in the Department for the pathological anatomy and the histopathology of the Military hospital in the Zagreb.
Investigated were 195 samples of the mucous membranes,
altogether 1.600 microscopic preparations.
Calculation of the throughput.
The circulation network was studied by a microscope. For each examined the segment of circulation (CA, CV, MA, a-v "by-pass", and others) was studied 50 villuses in which the same was best presented. There were measured lengths and widths of villuses,
diameter of central lacteals, CA, CV, and-v "by-passes" on the top, capillaries, a-v "by-passes" in the middle, MA and a-v "by-passes" in the region of crypts. Lengths were calculated towards Lewit’s rabbit (with proportion and conversion’s factor). Obtained results were checked under the microscope. From lengths and radius were calculated SR (segmental resistance) and carried out necessary conclusions.
From the length and widths of villuses were calculated lengths others tested elements.
Lengths tested elements of the Lewit’s rabbit:
CA 500,0 µm – conversion factor towards the length of the villuses
l,0.
CV 500,0 µm - conversion factor towards the length of the villuses 1,0.
A-V "b-p" on the top of villuses 180,0 µm – conversion factor
towards the length of the villuses 1,4.
Capillaries 80,0 µm - conversion factor towards the length of the
villuses 0,16.
A-V "b-p" in the middle 100,0 µm – conversion factor towards the
length of the villuses 0,76.
MA 100,0 µm - conversion factor towards the length of the villuses
0,2.
A-V "b-p" crypts - conversion factor towards the length of the
villuses 0,9.
After calculation of lengths, by help of the relationships SR = l/r on 4, were calculated segmental resistances of tested segments of the circulation and carried out necessary conclusions.
From tables of frequencies were calculated indexes and percentages of decrease or magnifications of all investigated variables.
The throughput was calculated as follows: Volumes of vilues were calculated by formula for the roller: V = r² . 3,14 . v Number of villuses in the gram of tissues was calculated by
proportion towards the volume numbers of villuses at Lewit’s rabbit.
The throughput through the gram of the mucosa before the vagotomy is constant and is 1,18 ml/min/g.
The throughput through the gram of the intestinuma before vagotomy is constant and is 1,242 ml/min/g of tissues.
The through through one villus was calculated dividing of the mucous throughput through the gram of tissue with the number of villuses in one gram of tissue.
The throughput through the region of crypts of one villuses was calculated dividing the mucous throughput through the region of crypts with number of villuses in the gram of tissues.
The throughput through the region of villuses of one villuses was calculated dividing the mucous throughput through the region of villuses with the number of villuses in the gram of tissues.
The throughput through the region of crypts, before vagotomy, is 44 %, and through the region of villuses 56 % of the throughput through the region of villuses.
The throughput after vagotomy was calculated by proportion towards earlier presented results of Lewitt’s work:The radius of CA is decreased for 30 % what reduces the mucous throughput with 1,l8 ml/min on 0,81 ml/min/g. In the same time the throughput through the region of crypts grows with 44 % on 60 % of the total mucous throughputs. For the calculation of the throughput was used following relationships:
r/x 31,4
/100 – (---)on 2 . 100/. -----
r/o 51
Q/xm = (--------------------------------) • Q/o
100
Q/ xm = the mucous throughput in ml/min/g of of tissues in the determined time’s interval after vagotomy.
Q/o = the mucosa throughput through one gram of tissue in ml/min/g before vagotomy /1,18/.
r/x = radius of CA in the determined time’s interval after vagotomy.
r/o = radius of CA before vagotomy.
31,4 = the fall of the throughput in percentages at taper of CA for 30 % (more correct 31,36).
51,0 = percentage’s decrease of the cross-section of CA at decrease of radius for 30 %.
Qxk = (44 + 16/31,36 • x) • Qxm/100
Qxk = the throughput through the region of crypts in the gram of tissues in the determined time’s interval after vagotomy.
44 = the percentage of the throughput through the region of crypts before vagotomy.
16 = the increase of the throughput in percentages through the region of crypts at decrease of the mucous throughput for 31,36 %.
x = the percentage of reduction (or increases) the mucous throughput through the gram of tissue determined time’s interval after vagotomy.
Qxv = (56 – 18/31,36x) • Qxm/100
Qxv = the mucous throughput through the region of villuses in the determined time’s interval after vagotomy. 56 = the percentage of the throughput through the region of villuses before vagotomy.
18 = decrease of the throughput in percentages through the region of villuses at decrease of the mucous throughput for 31,36 %.
Qxi =(Qxm/19) • 20
Qxi = the intestinal throughput through the gram of tissues.
From tables of frequencies was carried out the statistical processing of results. There were calculated the arithmetic midpoints and measured variability. Determined is P = the statistical probability, as measures of a chance of appearing.
III. RESULTS
The amount of dates does not allow theirs complete presentation. Therefore, we decided to present only variables and theirs arithmetic’s midpoints, which are indispensable for understanding of the work, and carried out of the statistical batch processing. That is tables of frequencies of arithmetic features, and variability of measured characteristics, and verisimilitudes as a chance of appearing determined reactions.
How it is going for the verisimilitude system (not determined) were used statistical methods. Hypothesis of the work is reductions of circulation after vagotomy. Reducing circulation and its redistribution is a happening that not have to happen obligatory (stochastic correlation). Verisimilitude, as for this the most appropriate statistical method, measures a chance of appearing, and has a value from 0 up to 1. (1 = happening must appears; 0 = impossible event.)
m
P = ---
n
P = verisimilitude
m = number of favorable outcomes
n = number of total possible outcomes
Variability of measured characteristics features is presented in tables of variability of the mucous throughput in ml/min/g of tissue. Variability of others features is not presented, although it is discussed in tables of the arithmetic midpoints (in the supplement). Variability of the radius of central arterioles is in the linear correlation with variability of the mucous throughput in ml/min/g of tissue, so that a special presentation is not needed.
In all tested cases appeared a strong reduction and redistribution of circulation, so P = 1, what means that hypothetical set event must appears. VARIABILITY OF THE MUCOUS THROUGHPUT in ml/min/g of tissue:
I. 1 3 10 20 30 45 60 90
D +0,747 +0,309 -0,101 -0,094 -0,221 -0,089 +0,298 -0223
J +0,18 -0,03 -0,193 -0,197 -0,142 -0,086 +0,085 -0,050
etc.
PROBABILITY
m
P = ---
n
P = probability
n = number of total possible outcomes
m = number of favorable outcomes
P 1 3 10 20 30 45 60 90
D 0,7 1,0 1,0 1,0 1,0 0,9 0,9 1,0
J 0,6 1,0 1,0 1,0 0,9 1,0 1,0 1,0
m = 8
n = 8
P = 1 – happening must appears.
System becomes determined. Satisfies mathematical processing of dates.
ANALYSIS OF DATES
1 to 3 minutes after vagotomy duodenal villuses lengthened with 421,56 on 523,09 µm (24,08 %). Width of villuses increases with 130,50 on 141,30 µm (8,28 %). Increased also the radius of the central lakteal with 8,0 on 8,37 µm (4,62 %}. Radius a-v "b-p" on the top increased with 4,41 on 4,83 µm (9,52 %), capillaries reduced with 1,69 on 1,65 µm (2,37 %), CA with 2,65 on 1,59 µm (40 %), CV with 4,13 on 3,48 µm (15,74 %), MA with 2,02 on 1,99 µm (1,49 %). Radius a-v "b-p" in the middle increased with 2,55 on 2,59 µm (1,57 %}, and in the region of crypts with 6,70 on 6,93 µm (3,4 %).
Changed also SR (segmental resistance), and it: CA with 48,91 on 198,93, CV with 3,l6 on 5,76, "b-p" on the top with 3,57 on 0,44, MA with 4,76 on 8,72 and capillaries with 7,87 on 13,05. Segmental resistance a-v "b-p" in the middle reduced with 5,84 on 3,27. In the first three minutes after vagotomy happened importance redistributions of the throughput. Contraction of pre-capillary arterioles reduced the throughput through capillary areas. SR of CA grows for 306,73 %, and CV only for 82,28 %. SR "b-p" on the reduced for 99,88 %. How the capillary resistance grew for 65,82 %, majority of blood that in the CA arrives directly from the big vessels in submucosas increasingly directs with "b-p" on the top in CV. SR MA grew for 83,19 % (CA for 306,73 %), while a-v "b-p" in the middle was lowered with 5,84 on 3,27 (44,,01 %). Through this "b-p" and marginal arterioles directs relative larger throughput than through the top of the villuses. Vagotomy reduced the total mucous throughput, intensifying at the same time the throughput through all "b-p" -es.
A-v "b-p"–es in the area of crypts opened with the diameter of 13,86 µm. (Before of vagotomy 1 µm.) Majority of the circulative blood, 1 to 3 minutes after vagotomy, directed through this connection (0,437 ml/min/g). The throughput through the region of crypts was so puffed up on the account of the throughput through villuses (0,224 ml/min/g). Secretive capillaries are turning into resorptive. Organism adapted on the hypovolemia, resorbing s and draining the intestinal liquid via "b-p" in the middle of villuses. Reaction is identical that in the hypovolemic shock.
Redistribution of the throughput through the mucosa of jejunums 3 minutes after vagotomy is similar to duodenal. Lengths of villuses reduced with 579.18, before vagotomy, on 528,82 µm after. Width of the villuses increased with 1l3,94 in the first minute on 140,44 µm in the third. Radius CL is increased on 8,85 µm (6,97 before the vagotomy, and 7,38 one minute after). Radiuses CA (3,82 - 2,73 – 3,18}, "b-p" of tops (5,41 - 5,33 – 5,02), capillaries (1,98 - 1,82 – 1,38), "b-p" of the middle of the villuses (2,97 – 2,70 -2,25), MA (2,23 - 2,05 – 1,83). Radius CV in the first minute is reduced with 5,05 on 4,19, in order to in third would have increased on 5,94 µm. "B-p"-es in the region pf crypts were widen opened with the diameter of 16,88 µm in the first minute, and 17,88 µm in the third.
The segmental resistance of CA, in relation to that before vagotomy increased 46,95 times, CV 13,1 times, "b-p" on the top 4.39 times, in the middle 3,5 times, capillaries 4.32 times. Here is also the throughput through all "b-p"-es accelerated. How SR capillaries were increased only 4.3 times, and MA 1,8 times, we concluded that vagotomy guards the circulation of the basal layers of villuses. Secretive capillaries were turning into absorptive.
1 to 3 minutes after vagotomy carried out a strong reduction and redistribution of the throughput. Blood was increasingly directed through a-v "b-p"-es. The largest part went via "b -p"-es in the region of crypts, whose diameter increased with 0,01 µm before vagotomy on 17,88 µm in the third after vagotomy. It is useful to mention that SR of these "b-p"-es was only 0,06 in the first minute and 0.035 in third. Progressively declining of SR on the direction of CA - "b-p" on the top, CV, as well as the relative growth of SR of capillaries and pre-capillary arterioles undoubtedly points out the strong redistribution of the reduced throughput in benefit of deeper layers.
In the third minutes after vagotomy the throughput through the mucosa of jejunum was 0,627 ml/min/g of tissues, of which through the region of crypts 0,418, and vilusa 0,190. The throughput through the mucosa reduced for 46,98 %, of which through the region of crypts 19,67 %, and through the region of villuses (most superficial layers) for 71,19 %.
In 10th minutes after vagotomy changes of decrease and redistributions of flow are continuing. Radius CA duodenal villuses reduced on 1,48 µm, "b-p" on top on 4,15 µm, CV on 2,79 µm, capillaries on 1,22 µm, a-v "b-p" in the middle on 1,98, MA on 1,53 µm. A-v "b-p" in the region of crypts was still open (radius 7,39). SR of CA grows with 6,88 on 284,66 (41,37 times), CV with 1,47 on 83,54 (56,83 times); and-v "b-p" on the top with 0,34 on 56,91 (167,38 times), and-v "b-p" in the middle with 4,83 on 1200,86 (248,63 times), MA with 5,68 on 192,80 (33,94 times) and capillaries with 4,53 on 98,63 (21,38 times).
The smallest growth of the relative segmental resistances we had at capillaries (21,38 times), then at MA (33,94 times). Follow CA (41,37 times), CV (56,83 times}, "b-p" top (167,38 times) and "b-p" in the middle (1200,86 times). As the resistance in capillaries grew only for 21,38 times, and SR "b-p" in the middle of villuses for 1200,86 times, the blood beside progressive strong reduction of the throughputs was kept in capillaries. Organism resisted to emptying of capillary spaces. This conclusion is a result of analyses of the arithmetic midpoints. Organism, namely, in particular cases really resisted to emptying of capillary spaces with strong contraction of drained vessels. In our material, it happened only once, then, when reduction of circulation was the largest. In all rest cases SR "b-p" in the middle of villuses progressively declined, with 4,83 before the vagotomy, on 10,56 in 10th minute after it. Drainage of the blood hereby progressively increased. So was increased the absorptive function of capillaries. (Were reduced the hydrostatic and oncotic pressure.) Only in one case, SR of the draining a-v “b-p” in the middle of the villus grew insomuch average that the average for all experimental animals gave different results. If we compare others sizes (in length and widths of villuses with diameter CL, radiuses arteries we will see that in that case (the first experimental animal, the rat, not rabbit, tested for comparing) achieved bigger reductions of all elements (rabbits), with it the throughput too. The average throughput of the mucosa of duodenum in 10th minutes after vagotomy is 0,634 ml/min/g of tissue, and at the first experimental animal (the rat) 0,533. The throughput via the most superficial layers was 0,197 and 0,131. In the villus 1,04 x 10 on the minus 5, and 4,31 x 10 on the minus 6. Such strong reductions of flows through the most superficial layers caused also appropriate reaction of organisms. Conclusion: Vagotomy beside a strong reduction and redistribution of circulation guards elementary functions of intestinal uvulas.
Similar changes were also in the jejunal mucosa. Lengths of villuses were reduced with 579,18 on 490,20 µm (15,36 %).
The diameter of villuses were increased with 128,65 on 142,28 µm, and of CL with l3,94 on 17,10 µm.
Radius a-v "b-p"on the top reduced with 5,41 on 3,53 µm (34,75 %), capillaries with 1,98 on 1,40 µm (29,29 %), CA with 3,82 on 1,83 (52,09 %), CV with 5,05 on 2,60 (51,49 %), MA with 2,23 on 1,76 (21,08 %), a-v "b-p” in the middle with 2,97 on 2,34 (21,21 %). Radius a-v "b-p" in the region of crypts reduced with 8,94 the mm in the third minute after vagotomy on 8,34 µm in tenth, what is a result more strongly reduction of circulation through all layers.
SR of CA of jejunalnog vilusa grows, towards the condition before the vagotomy, for 150,3 times, CV for 95,97 times, "b-p" on top for 7,l8 times, "b-p" in the middle 65,69 times, MA for 91,31 times, capillaries 55,97 times. Changes of SR suggest keeping back the blood in capillaries and "b-p"-es on the top of villusses.
There was reduced also the throughput of pre-mucous parts. Because of the strong contraction of smooth musculature was reduced the total volume of the intestine.
20 minutes after vagotomy changes of tested elements of circulation of the duodenal mucosa suggest additional reduction of circulation. Radius "b-p" on the top of villuses was 3,59 µm (before vagotomy 5,41, in 10 minutes after 3,53), CA 1,54 (3,82 and 1,83), CV 2,5l (5,05 and 2,66), capillares 1,44 (1,98 and 1,40), "b-p" in the middle 2,29 (2,97 and 2,25), MA 1,69 (2,23 and 1,76), "b-p" in the region of crypts 8,34 (0,1 and 7,59.
There was changed also SR of tested segments of circulation. In the direction CA, "b-p" the top, CV (400,22 - 8,10 - 11,02) still ever we had relatively smaller resistance to the streaming of the blood through a-v anastomoses is than through CV. As SR MA 128,85, the blood was increasingly directed hereby and "b-p"-es in the middle of villuses whose SR was 68,91. The tops of villuses was contracted, and circulation reduced on the most basically functions.
The strong reduction of circulation was found also in the jejunum. Radius a-v "b-p" on the top of villuses was 3,59 µm (5,41 and 3,53), CA 1,54 (3,82 and 1,83), CV 2,51 (5,05 and 2,66), capillary 1,44 (1,98 and 1,40), and-v " b-p " in the middle 2,29 (2,97 and 2,34), MA 1,69 {2,23 and 1,76).
Beside tendency of returning some elements of circulation towards normal (gradually giving in spasms of the smooth musculature) the total throughput was still strongly reduced, the top of villuses were contracted, and circulation reduced on minimum. This was best seen from relative SR of tested segments of circulation. Sudden fall of SR in "b-p"-es in the middle of villuses enabled the fast flow of the reduced amounts of blood through main blood vessels with relatively better-preserved circulation of capillary areas. The throughput was still ever strongly reduced; radiuses CA and MA were still ever progressively reducing.
30 minutes after vagotomy circulation through mucosa of duodenum and jejunums was still ever strongly reduced, beside tendency for returning to normal.
Radius of CA duodenal vilusa was 1,69 (3,40 and the l.,55), CV 2,59 (4,78 and 2,65), MA 1,53 (2,15 and 1,67), and-v "b-p" on top 3,93 (5,31 and 4,2l), "b-p" in the middle 1,95 (2,90 and 2,ll), and-v "b-p" of the region of crypts 7,01 (0,0l and 7,67). SR CA of duodenal villuses were 14236,65 (6.88 and 400,22), CV 257,27 (1,47 and the LL,02) "b-p" on the top of villuses 41,13 (0,34 and 8,10), "b-p" in the middle 5306,02 (4,83 1 68,91), MA 195,43 (5,68 and 8,10), "b-p" in the middle 5306,02 (4,83 and 68,91), MA 195,43 (5,68 and 128,45) and capillaries 2266,46 (4,53 and 98,63). Average values neither here were, as earlier at "b-p" in the middle of villuses in 10th minutes after vagotomy, reflection of a real situation. At one number of experimental animals circulation was hurriedly returning to normal (In the fifths and the ninth experimental animals were 1,l1 and 1,0l ml/min/g of tissue with still ever strong reduction of circulation in the region of villuses (the most superficial layers). In six experimental animals still ever there were a progressive reduction of circulation, while in others two was noticed a mild growth towards the condition in twentieth minute after vagotomy. SR "b-p"-es in the region of crypts were only 0,6. Radius of CA jejunal villuses were 1,97 (3,82 and 1,54), CV 3,45 (5,05 and 2,51) MA 1,88 (2,23 and 1,69}, a-v "b-p"-es on the top 5.38 (5,41 and 3,59), a-v "b-p" in the middle 2,48 (2,97 and 2,29), a-v "b-p" in the region of crypts 9,72 (0,01 and 8,34). SR of tested segments of circulation of jejunal villuses were pointed at gradually returning the circulation on the condition before vagotomy, beside still a strong reduction of circulation through the most superficially layers. SR CA were 64,52 (5,19 and 563,13), CV 6,68 (1,0 and 78,31), "b-p"-es on the top (0,33 and 2,7), capillaries 24,73 (6,19 and 57,78), "b-p" in the middle 3,47 (1,63 and 30,86), MA 97,30 (12,38 and 69,10) and "b-p"-es in the region of crypts 0,l7.
45 minutes after vagotomy, contraction of the smooth musculatures of duodenal villuses yields. Jejunal villus was contrary of it strongly contracted. The mucous throughput of duodenum was 0,742 ml/min/g of tissue, and jejunums 0,710.
SR CA of duodenal villuses were 340,67 (6,88 and l4236,65), CV 69,56 (1,47 and 257,27), "b-p" above 1,09 (0,34 and 41,13), capillary 24,19 (4,53 and 2266,46), "b-p" in the middle 12,48 (4,83 and 5306,02), MA 26,48 (5,68 and 195,43), "b-p" crypts 2,99 (0,6 in the thirtieth minute after vagotomy). Radiuses CA were 1,83 (3,40 and 1,69), CV 2,74 (4,78 and 2,59), "b-p"-es at the top 4,35 (5,31 and 3,93), capillaries 1,36 (2,05 and 1,2l), "b-p"-es in the middle 2,21 (1,95 and 2,90), MA 1,62 (2,l5 and 1,53) and "b-p"-es in the region of crypts 8,42 (7,01 in the thirtieth minutes after vagotomy).
Similar changes happened also in the jejunal mucosa. Radiuses CA were 4,07 (3,82 and 1,97), CV 3,23 (5,05 and 3,45), "b-p" on the top 4,93 (5,41 and 5,38), capillaries 1,59 (1,98 and 1,48), "b-p the" in the middle of villuses 2,32 (2,97 and 2,48), MA l.,82 (2,23 and the l.,88) and "b-p" in the region of crypts 7,8 (0,01 and 9,72). SR CA were 106,94 (5,l9 and 64,52), CV 12,79 (1,0 and 6,68), "b-p" on the top 10,61 (0,33 and 0,55), capillaries l4,92 (6,19 and 24,73), "b-p" in the middle of villuses 5,04 (1,63 and 3,47, MA 8,65 (97,13 and]2,38), "b-p" in the region of crypts 0,05 (0,17).
Beside smaller oscillations the trend of returning to the condition before vagotomy was continuing. But, in spite of it the total flow was still strong reduced.
60 minutes after vagotomy the mucous throughput of duodenum was 0,812 (1,18 and 0742), from that through the region of crypts 0,463, and through the region of villuses 0,344 ml/min/g of tissue. Radius CA were 1,85 µm, CV 3,07 µm, "b-p" on the top 4,10 µm, capillaries 1,43 µm, "b-p" in the middle of villuses 2,l6 µm and MA 1,75 µm. SR CA were 384,73, CV 4,99, "b-p" on the top l.,08, capillaries 38,84, "b-p" in the middle 5,57, MA l0,l4 and "b-p" in the region of crypts 0,12. By oscillatory returning the normal circulation was increasingly faster repairing in the region of crypts than in the region of villuses (most superficial layers).
Similar changes were found in the jejunal mucosa too.
90 minutes after vagotomy, circulation was farther reducing: in the duodenum on 0,652 ml/min/g, and in the jejunum on 0,676 ml/min/g. This pointed at slow returning to normal, perhaps days long!
Initial post-vagotomic reactions of the mucous membranes of duodenum and jejunum
were examined in the first minute after vagotomy in 10 experimental animals. It was recorded the initially decrease of volume of the villuses because of contraction of the smooth musculature. This is importantly because volumes of villuses, in the period of the largest reduction of flow in 3, 10, 20 and 30 minutes after vagotomy were increased (in duodenum) or minimally reduces (in jejunum). A decrease of volumes of villuses was happening, namely, later in 45, 60 and 90 minutes after vagotomy, when circulation was recovering. It was noticed diminishing of radiuses of all tested elements of circulation. So radiuses CA of the duodenal villuses were average 2,65 µm (before vagotomy 3,40), CV 4,13 (4,78), "b-p" on the top 4,41 (5,31), capillaries 1,69 (2,05), "b-p" in the middle 2,55 (2,90), MA 2,02 (2,15). "B-p"-es in the region of crypts were widen opened and theirs radiuses were 6,70 µm. Only radius "b-p" on the tops were slightly increased, what was a result of simultaneous contracting CA and CV.
Presented results are arithmetic middle values of tested segments of circulation all experimental animals. They, however, do not give a correct display of the initial reaction. During analysis of the verisimilitudes of reductions of circulation, we found that in the first minute after vagotomy in the duodenum they were 0,7, and in the jejunum 0,6, while is total P l,0. In the first minute on the vagotomy in 3 cases in 10th minute after vagotomy in duodenum was not obtained expected reduction of circulation. In the jejunum this was happened in 4 animals. Reaction is not, therefore, probable and obligatory. Results demand the statistical processing.
Probability as a measures of coincidences (the chance of appearing), will present us to dichotomy of measured characteristics.
P CA CV b-p top capilaries b-p middle MA b-p crypt
D 0,7 0,6 0,7 0,8 0,8 0,5 0,7
J 0,8 0,6 0,5 0,7 0,4 0,6 1,0
_____________________________________________________________
0,75 0,6 0,6 0,75 0,6 0,55 0,85
From obtained values is visibly the order of contractions of tested segments of circulation, and with it also reductions of it, which are disharmonious. The first were contracted CA and capillaries, then CV, "b-p"-es on the top and "b-p"-es in the middle. Circulation in the region of crypts was intensified, as well as in basal parts of villuses (MA and "b-p"-es in the region of crypts). Reduction of circulation appeared in 7 cases in duodenum and in 6 in jejunum (P 0,7 and 0,6). But, in spite of it total P is 1. The strong reduction of circulation was achieved, namely, only in the third minute after vagotomy.
Influence of a hypoxia on the post vagotomic circulation of duodenal and jejunal mucous membranes, and vice versa,
was examined in four experimental animals (B, C, D and E). Hypoxia achieved with the passage of the vascular network with gelatin of the same viscosity with blood during several minutes. Samples of the mucous membrane was taken from 10 to 30 minutes after vagotomy. In the hypoxic intestine was confirmed a contractile post-vagotomic reaction of smooth musculature of villuses and blood vessels. In the previous shown series of 10 experimental animals was achieves the largest decrease of radiuses CA of 54,4l % in duodenum and 59,69 % in jejunum. Radiuses of CA of hypoxic duodenum were reduced for 70,59 %, and jejunum for 53,66 %. In accordance with it were changed SR of blood vessels and the throughput. The maximally decrease of the throughput in duodenum was 56,69 %, what is 0,511 ml/min/g of tissue. The throughput through the region of crypts was reduced, at the same time, for 38,32 %, and in the region of villuses for 81,69 %. In the previous series of 10 experimental animals the throughput was 0,628 mil/min/g of tissue (decrease 46,74 %), while the throughput through the region of crypts was 0,424 mil/min/g (decrease l8,57 %), and through the region of villuses 0,l86 mil/min/g (decrease 71,95 %). The maximally decrease of the throughput of the hypoxic jejunum was 45,93 % (0,638 mil/min/g). The throughput through the region of crypts, at the same time, was decreased for 54,07 % and was out 0,414 mil/min/g of tissue, and through the region of villuses 68,53% and was 0,208 mil/min/g. In the previous series of 10 experimental animals results were: 0,627, 0,418 and 0,190 ml/min/g in third minute and 0,659, 0,416 and 0,227 in the twentieth minute after vagotomy. Decrease in percentages in the third minute after vagotomy was 46,98 %, 19,67 % and 71,l9 %, and in the twentieth 44,28, 21,67 and 65,60 %. Hypothetical reductions of the throughput came in both series, in all 14 experimental animals. (P = 1) Conclusion: A moderated hypoxia does not change significantly post-vagotomic reaction of blood vessels.
Influence of a hypercapnia on the post-vagotomic reaction of blood vessels of duodenum and jejunums
was examined in two experimental animals. There was used the device for the extracorporeal circulation. Hypercapnia was achieved with passage of gelatin of the same viscosity with blood during 15 to 20 minutes. Radius CA of the duodenal villuses increased for 81,47 %, and the total mucous throughput for 139,83 %. Radius CA of the jejunal villuses increased for ll5,97 %, and the throughput for 88,14 %. The mucous throughput of duodenum was 2,83 mil/min/g and jejunum 2,22 ml/min/g. Conclusion: In hypercapnia post vagotomic contracted blood vessels of the mucous membranes of duodenum and jejunum became wider, and the total throughput through duodenum and jejunum increased. Cause for it is likely a paralysis of vasomotor nerves.
IV. DISCUSSION
Post-vagotomic contractile reactions of mucoses of duodenum and jejunum were caused by contraction of the muscular threads. Because of it, villuses became smaller and fatter, in the total volume smaller. Spaces among villuses became widened, openings of crypts larger. Mikrovili of endocrine cells of GEP system more easily receive information from the lumen on based of which they discharge theirs granules in peri-capillary spaces of the sub-epithelial capillary layer, mostly basal parts of villuses and the region of crypts. In this way, indirectly, these cells post-vagotomic influence on the mucous circulation. But, it is not the subject of this work.
There was also eased emptying of Brunner glands. (Vagotomy reduces total secretion of all glands.)
Contraction breadthways helps the network of elastic fibers, which penetrate in villuses and in the area of crypts directly from smooth muscular fibers. With contraction of the smooth musculature contract also the network of these fibers, and so abridge the region of crypts and villuses longwise and breadthways. The smooth musculature of the mucosa layer has two layers, inside mostly circular and outer mostly longitudinal.
We said that a-v anastomose on the top of villuses important for the heart functioning, because in a non digestive phase blood directly, over this anastomose goes in veins, not spending energy for pushing blood through capillaries. For the heart are important also all others anastomoses, which after vagotomy redistribute blood from superficial in deeper layers. Important are also cells of GEP system, which over MA and sub-epithelial capillary layer of basal parts of villuses influence on circulation of a-v connections in the middle of villuses. .
Maximal contractions of the smooth muscular fibers of the mucous membrane shrinks and shortens the top of villuses contracting "b-p"-es on the top with a pressure from outside.
Through the region of crypts passes large central arterioles of villuses, which start from sub-mucous veins, penetrate the muscu1aris mucosa, enter in the region of crypts, and go parallel with central hiluses towards the top of villuses without branching. That we also said for smaller branches of sub-mucous arteries, which after penetrating the muskularis mucosa, enter in this region. From them emerge MA and the sub-epithelial capillary layer of the region of crypts. Before of vagotomy blood streams through these vessels passing by a-v anastomoses this region. Opening and exists anastomoses is visibly also in microscopic preparations obtained 3, 10, 20, 30, 45, 60 and 90 minutes after vagotomy. For a correct differentiating of these connections is necessary additional investigating.
Vagotomy influences also on the lymph circulation of the mucous membrane of the small intestine. We already said that absorbed liquid gathers in the interstitial tissue, and that from here go away via blood and lymph capillaries. The hydrostatic and the oncotic pressures determine quantity of taken away liquid with every of these two systems. With accumulation of liquid grows the interstitial volume and the hydrostatic pressure, while the onkotski pressure falls. How the interstitial tissue is composed from collagens’ fibers and mucopolysaccharides, which are mutually interwoven in the gelatinous structure, hydraulic conductivity of normally hydrated interstitial tissue is very small. How the volume conductivity is vice versa proportional to the concentration of the chialuronic acid, the grows of the volume of liquid increases the hydraulic conductivity of the intestinal tissue. The oncotic pressure of the interstitial liquid falls. Also stops the capillary filtration. Filterable capillaries turn into absorptive. Lymph vessels fill also with liquid. Divisions of the mucous interstitial volume on central and the juxtacapilary parts ensures the passive flow of liquid in capillaries. The hydrostatic pressure is in juxtacapillaries’ areas more sensitive at changes of the volume of that in central interstitial spaces. Therefore a larger part of absorbed liquid takes away hereby. By an accumulation of liquid the largest part of the fall of oncotic pressure is visible just in that fragmentary sub-epithelial-juxtacapillary layer.
By post-vagotomic contraction of the smooth musculature of villuses and blood vessels, grow the hydrostatic pressure, and become faster the passive flow of liquid in blood and lymph capillaries; grows also the colloidal-osmotic pressure of the interstitial fluid and concentration of the hyaluronic acid. Secretive capillaries become absorptive. Because of it, the contractive reaction of capillaries is less prompt from others segments of circulation. Larger part of liquid is taken away hereby directly in circulation. When colloidal-osmotic pressure grows, grows also concentration of the hyaluronic acid. The hydraulic conductivity falls, and liquid passively no longer penetrates in lymph vessels. The hydrostatic pressure pushes remaining liquid in the blood vessels. Because of the contraction of pre-capillary arterioles, the pressure in capillaries falls.
It would be interesting to examine the role of GEP cells on circulation after vagotomy. Some gastro-intestinal hormones (cholecystokinin and the secretin) increase the intestinal flow of blood and lymph if they are instilled in a. mesenterica crania1is. Turner and Barrowman suggest that hormones exude during absorptions cause vascular changes (increased capillary perfusion and capillary pressure).
The absorbed liquid increases the capillary’s hydrostatic pressure. As the hydraulic conductivity of the intestinal tissue reduced, because of considerable increase of concentration of the hyaluronic acid and increased the oncotic pressure, the capillary filling of sub-epithelial capillary’s layer increasingly carry's out with absorption of liquid from the intestinal lumen, and all more less from interstitial tissue. Organism compensates weaker perfusion with increased absorption of capillaries.
In an absorptive phase of animals, which are not subjected to vagotomy, absorbed liquid spreads the interstitial space, grows the interstitial hydrostatic pressure and reduces oncotic. Changes cause flow of liquid from interstitial tissue in blood.
After vagotomy the interstitial space becomes
narrower and the hydrostatic and the oncotic pressures grow; increases also concentrations of the hyaluronic acid, and pre-capillary arterioles contract. The oncotic pressure in capillaries falls, because of absorption of liquid. In later phase increasingly less liquids is absorbed from interstitial tissue, more, and more, from the sub-epithelial juxta-capillary lymph space, immediately after absorption from the intestine.
Weaker contractions of capillaries can be explained with the role of sympathicus. In the introduction we said that noradrenergic fibers go next to blood vessels and that are, especially well nerved the muskularis mucosa and the sub-mucosa. Dense solar plexuses surround blood vessels, especially in the region of crypts and in the sub-epithelial capillary layer of villuses and base-lateral membrane. The final sympathetic fibers of these plexuses terminate in capillaries. Contraction seems to be directly proportional to mass of the smooth musculature of appropriate segment of circulation. It seems that all is regulated so.
Sympathicus plays the crucial role in the protection of the absorbed function of the intestine after vagotomy. Relatively weak the smooth musculatures of capillary veins make easier this function. Levens and coworkers think that alpha-receptors on the base-lateral membrane of epithelial cells bind noradrenalin on nerve final threads and accelerate transport of fluid. Wit this role of sympathicus is able interpret also relative slower and weaker contraction, not just capillaries, than also connections of these with varicosities nearby base-lateral membranes.
Vagotomy, as well as stimulation of noradrenergic fibers of villuses, reduces lymph flow, coefficient of the capillary filtrations and pressure. It also reduces the pressure of the interstitial circulations and there is no so called "self-regulation". It was said that stimulation of sympathetic fibers of the small intestine promptly reduces the blood throughput, which after it gradually gives back in normal values. Vagotomy, on the other hand, initially, promptly poorly reduces the blood throughput. We can freely to say that it makes it something slower and initial oscillatory. Reaction is the strongest in the first 30 minutes after vagotomy. Actively are contracting arterioles and venulas and it balanced, arterioles something weaker of venulas. Because of this biphasic answer on a vagotomy, the capillary pressure does not depreciate considerably. Beside this rapid fall of the intestinal mucous throughput, capillaries are still ever weaker contracted of arterioles and venulas. However, they are in this phase still ever secretive although
gradually grow their absorptive function.
After adrenergic stimulation the intestinal throughput inside the first 5 minutes falls on 48,3 % with oscillations about 7 %, in order later be increased on 73,7 % with oscillations of 4,2 %. Coefficient of the capillary filtration of the adrenergic stimulated intestinal tissue is reduced on 75 % (later 25 %). After vagotomy reduction of the intestinal throughput, specially mucosal, is stronger. 10 minutes after vagotomy radius CA is 1,48 µm in the duodenum and 1,83 µm in the jejunum; 20 minutes after is 1,55 and 1,54; 30 minutes after is 1,69 and 1,97; 45 minutes is 1,83 and 2,07. In 60 minutes is 1,85 and 2,5. SR of CA of the duodenum grows with 6,88 before the vagotomy on 198,93 in the third minutes; 284,66 in tenth, 400,22 in twentieth and 14236,65 in thirtieth, when suddenly reduces that in 45 minutes is only 340,57. In the 60th minutes is again something bigger and is 384,73, and in ninetieth falls on 83,62. In the jejunum we have an increase from 5,19 before vagotomy on 243,68 in the third minute and 758,86 in tenth. In twentieth minute SR is 563,13, in thirtieth 64,52. In the 45th minute SR of CA of jejunal villuses is 106,94, in sixtieth 26,46, and in ninetieth 79,22.
At SR we have also a oscillatory returning to normal. SR of CV grows more slowly, and of "b-p" on top lesser than of CA and CV. (See tables of arithmetic midpoints!). The total interstitial resistance grows mostly at account of pre-capillary resistance, as well as the venous segmental resistance. Biphasic answer becomes increasingly significant.
Hereinto needs still ones look back at circulation in the region of crypts and the existence of numerous a-v connections in this area, which earlier are not described. Before a vagotomy a-v "b-p"-es of the region of crypts is not possible to notice, because they are contracted. 3, 10, 20, 30, 45, 60 and 90 minutes after vagotomy "b-p"-es of this region are dilated and the throughput is directed hereby by the most superficial layers of the duodenal and jejunal mucosa.
The theoretical model of the vascular network of mucosa of the small intestine, which in 1979 years made Lewite and coworkers, do not mention a-v "b-p"-es in the region of crypts. Neither others authors notice anywhere importance of these anastomoses, which only post-vagotomic become visible. Thanking to them the vascular network of regions of crypts and villuses make one functional
totality - vice versa of what so far was thinking. Lewite and coworkers are, namely, with injecting of the vascular network of the small intestine of rabbits with silicone’s foam and polystyrene’s micro-granules proved an independence of the blood throughput in the area of crypts from the throughput through villuses, what was completely understandable because those are two areas with different functions. With analysis of ours, microscopic preparations was visibly that it is correctly only in conditions in which the adrenergic system is not irritated, more precisely said, before vagotomy. In fact, after vagotomy circulation in the region of crypts and villuses is unique. A-v "b-p"-es in the region of crypts are turning aside the throughput by villuses.
The biggest fall of the mukoznog throughput is noticing between tenth and thirtieth minutes after vagotomy. Vagotomy redistributes the throughput in the direction of the regions of crypts.
From a special interest is analyses of the throughput in segments: capillaries, MA, and "b-p" in the middle of villuses. Contraction of capillaries lags after contraction of CA, CV and "b-p"-es on the top. Similarly is also with MA and "b-p"-es in the middle of villuses. Vagotomy guards the absorption function of the small intestine. Most poorly are contracted, and most quickly give back to normal "b -p"-es in the middle of villuses with which absorbed liquid directly takes away in the blood vessels. The throughput through the basal parts of villuses is relatively less reduced of that in apical parts.
Wit analysis of SR of these vessels we get still better insight in reduction and redistribution of the throughput, as well as in the role of "b-p"-es in the middle of villuses. (See tables of arithmetic midpoints!)
After vagotomy organism guards the most superficially layers from dehydration, maximally decreasing, simultaneously, circulation of them.
V. CONCLUSION
Vagotomy causes contraction of the smooth musculature of the intestine and reduces total the mucous and the pre-mucous throughput. Reduction is less prompt than after a sympathetic stimulation. The biggest decrease of the throughput achieves between 10th and 20th minutes after vagotomy, after it follows an oscillatory returning to normal, which lasts several hours, perhaps even weeks or months. With redistribution of the throughput, the blood is directed via the region of crypts. The biggest reduction of the throughput is in the most superficial layers (the region of villuses). The total mucous throughput reduces from 46,74 % in the duodenum on 46,98 % in the jejunum. Reduction of the throughput through the region of crypts is 20,92 % in the duodenum and 21,67 % in the jejunum. The throughput through the region of villuses reduces for 71,95 % in the duodenum and 71,19 % in the jejunum.
With vagotomy is probably, in certain cases, to stop bleeding from the superficial layers of the mucosa of duodenum and jejunum.
VI. COMPENDIUM - SUMMARY
1. In 10 experimental animals were examined the vascular answer of the duodenal and the jejunal mucous membranes 3, 10, 20, 30, 45, 60 and 90 minutes after vagotomy.
2. In 4 rabbits was examined the vascular answer the hypoxic duodenal and jejunal mucous membranes after vagotomy.
3. In 10 experimental animals were examined the initial post-vagotomic reaction of the duodenal and jejunal mucous membranes.
4. In 2 rabbits was examined the influence of vagotomy on the presence of a hypercapnia in the duodenal and the jejunal mucous membranes.
The segmental resistance of the vascular network is calculated by help of Poiseuille' law. Lengths and widths of villuses and radiuses of CL and others tested segments of circulation are measured under the microscope, analyzing 50 typical examples in the every sample. The total mucous throughput is got as a relationship towards Lewit’s rabbit. In the same way is calculated the redistribution of the throughput between the region of crypts and villuses. The hypoxic and the hypercapnic circulation is studied with use a device for the extracorporeal circulation and passage an isolated winding of the intestine with to the gelatin that is of the same viscosity with the blood. It is confirmed the hypothesis of the work: Vagotomy also reduces the mucous throughput of the duodenum and the jujunum. The biggest decrease was achieved between third and twentieth minutes after vagotomy, and it for 46,74 % in the duodenum and 46,98 % in the jejunum (in the hypocsic winding 56,69 and 45,93 %). The throughput through the region of crypts decreased for 20,92 % in the duodenum and 2l,67 % in the jejunum (in hypoxia 38,32 and 54,07); in the region of villuses for 71,95 % in the duodenum and 71,19 % in the jejunum (in hypoxia 81,69 and 68,53 %). After 30 minutes follows an oscillatory returning to normal.
Vagotomy is not in condition to reduce vasodilatation in hypercapnia. The total mucous throughput of a hypercapnic mucose after a vagotomy was increased for 139,83 % in the duodenum and 88,14 % in the jejunum.
There were described a-v anastomoses in the region of crypts. It was proved that is the post-vagotomic circulations of the region of crypts and villuses unique. The order and promptness of studied changes indicates that the post-vagotomic reactions dependent of mass of the smooth musculatures. Contractive reaction is
directly proportional to mass of the smooth musculatures.
Dissertation was made in the Departments for the Physiology, the Histology and the Pharmacology of the Faculty of Medicine in Zagreb, the Department for the Pathology of the Military hospital Zagreb, and in the Administrative and Technical Service of the Self-governing Interesting Community of the health care and the health insurance of the municipality of the Long Village (Dugo Selo).
Manager of work:
Prof. dr. sc. Mladen Stulhofer
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of rabbit small intestine.
Am. J. Physio1., 1979. Oct., 237(4):E 373-82.
13. Granger, D.N.,: Intestinal microcirculation and
transmucosal fluid transport,.
Am. J. Physiol., l98l. May, 240(5):G 343-9 (30 ref).
14. Granger, D.N. et al.: Intestinal blood flow.
Gastroenterology, 1980. Apr., 78(4):837-63 (316 ref).
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jejunal villus.
Acta Histochem. (Jena) 1983., 72(1):85-9
16. Shepherd, A.P.: Autoregulatory escape in the gut:
a systems analysis.
Gastroenterology, 65:77-91, 1973.
17. Granger, D.N. et al.: Sympathetic stimulation and
intestinal capillary fluid exchange.
Am. J. Physiol., 1984. Sep., 247(3 Pt 1):G 279-83.
18. Brunssen, I., et a1.: The effect of vasodilatation and
sympathetic nerve activation on net water absorption in
the cats small intestine.
Acta Physio1. Scand., 106:61-68. 1979.
19. Folkow, B.D.., et al: The effect of graded vasoconstrictor
fibre stimulation on the intestinal resi¬stance and
capacitance vessels.
Acta Physiol. Scand.: 6 1:445-457, 1964.
20. Lundgren, 0.: Role of splanchnic resistance vessels
in overall cardiovascular homeostasis.
Federation Proc. 42:1673-1677, 1983.
21. Greenway, C.V.: Neural control and autoregulatory
escape. In: Physiology of the Intestinal Circulation,
edited by A.P. Shepherd and D.N. Granger.
New York: Raven, 1984, chapt. 5 p 61-71.
22. Lundgren, 0., et al.: The effect of sympathetic
vaso¬onstrictor fibres on the distribution of capillary
flow in the intestine.
Acta Physiol. Scand.: 61:458-466, 1964.
23. Shepherd, A.P.: Intestinal 0-2 uptake during sympathetic
stimulation aud partial arterial occlusion.
Am. J. Physiol. 236 (Heart. Circ. Physiol. 15):
H 731-H 735, 1979.
24. Lundgren, 0. et al.: The effect of splanchnic nerve
stimulation on blood flow distribution, villous tissue¬
osmolarity and fluid and electrolyte transport in the
small. intestine of the cat.
Acta Physiol. Scand. 117:365, 1983.
25. Gidda, J..S. , et al.: Influence of vagus nerves on
electrical activity of opposum small intestine.
Am. J. Physiol., 1980. Nov., 239(51):G 406.
26. Lewit, D.G., et al.: Model for mucosal circulation of
rabbit small intestine.
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