Evidence for Reflex Control of Arterial Pressure from Abdominal

Evidence for Reflex Control of Arterial Pressurefrom Abdominal Receptors with Special

Reference to the PancreasBy .STANLEY J. SARXOFF, M.D., AND SAMUEL I. YAMADA, P H . D .

Hypotension in cat abdominal viscera, especially the pancreas, produced substantialelevations of arterial pressure and heart rate even in the presence of intact sinoaorticreceptors. Elevation of pressure in the distribution of the superior mesenteric arteryhad the reverse, effect. When viewed in the light of the neurophysiologie data ofGammon and Bronk, it would appear that these sites make a significant contributionto cardiovascular regulation in that species, presumably as a result of impulses arisingin abdominal pacinian corpuscles.

THE contribution of afferent impulsesfrom special sites in the vascular bed

to the regulation of arterial pressure has beenappreciated with increasing clarity1 since theobservations of Hering- in 1927. The mostimportant of these sites have, of course, beenthe carotid sinus and aortic arch. This com-munication describes experiments which sug-gest that abdominal pressoreeeptors may alsobe of importance.

METHOD

After preliminary experimentation with differ-ent anesthesias and various technics for producing

'reversible arterial occlusion, experiments wereperformed on 24 cats. The first 5 of these receivedehloralose-urethane (100 nig. and 1,000 mg./Kg.).The remaining 19 were given ehloralose alone (50-60 mg./Kg.). Additional anesthetic was rarelyrequired during the experiment. After making ainidline abdominal incision, silk ligatures wereplaced loosely around the three small arteries tothe head, body and tail of the pancreas (for pro-ducing pancreatic hypotension), around the coeliac,superior and inferior mesenteric afteries (for pro-ducing generalized visceral hypotension), aroundboth renal arteries, and around both commoncarotid arteries. The simultaneous interruption offlow in the coeliac, superior mesenteric and infe-rior mesenteric arteries will be referred to as thecombined vascular occlusion. An attempt wasmade to be as careful as possible to avoid injuryto any nerve structures in the vicinity of the arte-rial dissection. It was felt that these attempts

From the Laboratory of Cardiovascular Physiology,Xutiomil Heart Institute, Bethesila, M<1.

Received for publication November 4, 1958.

were more successful as additional experience wasobtained with the preparation. Kach periarterialligature was passed through thin polyethylenetubing, the end of which was slightly flared, sothat reversible occlusion could be readily pro-duced. Periods of occlusion were usually between1.5 and 2 min. Bilateral vagotomy was done eitherat the beginning or during the course of the ex-periment, after control observations had beenmade. Arterial pressure was measured in the fem-oral artery and occasionally also in a peripheralbranch of the superior mesenteric artery distal tothe point of occlusion of this vessel. In some ex-periments perfusion of the superior mesentericartery was carried out, after proximal occlusion,by repeatedly injecting through a catheter 1or 2 ml. of blood just withdrawn from a femoralartery catheter. Heart rate was recorded with aWater's cardiotachometer. All values were contin-uously recorded on a multichannel, direct-writingoscillograph. The effect of tctraethylammoniumchloride and hexanietlioniuiii chloride on the re-sponse to pancreatic and combined arterial occlu-sion was examined side by side with the effect ofthese agents on the response to bilateral commoncarotid artery occlusion. Heparinization, whenused, was accomplished by the intravenous injec-tion of 5 to 10 nil. of 1 per cent heparin in Kinger'ssolution. A heating pad under the animal servedto maintain body temperature. In 16 of the 24 eats,a total of between 5 and 30 ml. of 6 per cent dex-tran in isotonic saline was given intravenously overthe course of any given experiment when this wasrequired to maintain the control arterial pressurelevels.

The objective of these experiments was to deter-mine the reflex cardiovascular effects of changingthe pressure within the blood vessels of abdom-inal viscera, especially the pancreas.

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Fia. 1 Top left. Arterial pressure response to combined abdominal occlusion (coeliae, superiormesenteric and inferior mesenteric arteries) for 90 sec. Occlusion at first three signals;release at second three signals. Bilateral cervical vagotomy. Experiment 17. Top right.Comparison of pressor response to bilateral common carotid artery occlusion (C) and occlusionof three pancreatic arteries (JP) during periods designated by bars below C and P. Vagiintact. Experiment 8. Bottom. Arterial pressure response to combined occlusion for 10 min.Bar, 1 min. period. Vagi intact. Experiment 23. Ordinate, mm. Hg in all pressure tracingsfor this and subsequent figures. Three large squares, 1 min. in this and subsequent figures.

RESULTSThe temporary, combined occlusion of the

coeliae, superior and inferior mesenteric ar-teries (combined visceral hypotension) in thecat almost uniformly produced a brisk pres-sor response. Rarely these responses were aslittle as 20 mm. Hg., especially when the re-sponse to carotid artery occlusion was rela-tively brisk. More often they were in the vicin-ity of 85 to 100 mm. Hg. Arterial pressure in-crements of 125 mm. Hg or greater were notinfrequent (fig. 1 top left). The pressor re-sponse to pancreatic arterial occlusion alone(head, body and tail arteries) was generallytwo-thirds to three quarters of that observedwith the combined visceral artery occlusion(fig. 2). Pressor responses of as much as 100mm. Hg were observed during periods of pan-creatic vascular hypotension alone. Occasion-ally the response to the pancreatic and thecombined visceral hypotension were the same.

The time course, magnitude and contour ofthe pressor response to pancreatic hypotension

resembled that observed after common carotidartery occlusion in 2 eats (figs. 1 top rightand 4). In the other 22 cats, however, the re-sponse to either visceral or pancreatic hypo-tension was substantially greater than thatobserved after bilateral carotid occlusion.Figures 2 and 3 show the results of threesuch comparative experiments. The augmenta-tion of the pressor response to the combinedvascular occlusion by preliminary occlusionof both common carotid arteries is also shownin fiigure 3 Bottom.

In one cat before vagotomy the pressorresponses to the carotid artery occlusion, thecombined occlusion and the pancreatic arterialocclusion were 40, 25 and 20 mm. Hg re-spectively. After vagotomy the responseswere 95, 35 and 30 mm. Hg. After carotidsinus denervation the pressor response to thecombined occlusion increased to 100 mm. Hgand the response to pancreatic arterial occlu-sion rose to 50 mm. Hg. It appeared in thiscat, more emphatically than in the others,


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REFLEXES FROM ABDOMINAL RECEPTORS 327

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FIG. 2. Upper tracing, arterial pressure; lower tracing, heart rate in beats/min.; A, responseto bilateral common carotid artery occlusion. Signals mark occlusion and release. B, responseto pancreatic arterial occlusion; C, response to combined abdominal occlusion; D, response tobilateral renal artery occlusion; E, repeat of C; F, stepwise response to sequential occlusionof inferior mesenteric, superior mesenterie and coeliac arteries in that order at points indicatedby signals. Bilateral cervical vagotomy. Experiment 18. Pancreatic pacinian corpuscles ofthis cat are shown in figure 7.

that the abdominal pressoreceptors were heldin check by the sinoaortic sites.

Bilateral vagotomy did not adversely in-fluence the pressor response to either pan-creatic or general visceral hypotension. Con-trariwise, in those experiments in which acomparison was made immediately before andabout 10 min. after vagotomy, the responsewas intensified by vagus interruption (fig.3 Top).

Heart rate was continuously recorded in 17of the 24 experiments (figs. 2 and 3). The re-sponse to pancreatic or visceral hypotensionwas most often a tachycardia but this did notalways occur and in one instance a markedbradycardia was noted. With the vagi intactthe heart rate response to pancreatic vascularocclusion was tachycardia in 13, bradycardiain 1, and a mixed reaction with late brady-

cardia in 1 (15 experiments). The heart rateresponse to the combined visceral occlusionwas tachycardia in 8, bradycardia in 1 anda mixed reaction with late bradyeardia in 2(11 experiments) (fig. 3 Bottom left). Aftervagotomy was performed, tachycardia wasnoted in all 8 cats subjected to pancreatic vas-cular occlusion and in 3 of the 4 cats sub-jected to the combined vascular occlusion. Inthe fourth there was no change.

Of the 13 cats which exhibited tachycardiaduring pancreatic vascular occlusion with thevagi intact, 9 showed rate elevations whichwere greater than those seen during bilateralcommon carotid artery occlusion. Of the 8cats which showed tachycardia during thecombined vascular occlusion with the vagi in-tact, 6 showed rate elevations which weregreater than those seen during bilateral com-


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FIG. 3 Top. Response of arterial pressure and heart rate to bilateral common carotid arteryocclusion ( 4 ) , to pancreatic artery occlusion (13), and to pancreatic artery occlusion afterbilateral cervical vagotomy (C). Experiment 15. Left bottom. 'Response of arterial pressureand heart rate to combined abdominal occlusion. Sight Vottom. Response to bilateral carotidocclusion (long oar) and combined abdominal occlusion (short bar)» Vagi intact. Experiment 30.

moil carotid artery occlusion. After vagotomywas performed on the 8 cats in which tachy-cardia was observed during pancreatic vascularocclusion, 4 showed a greater elevation thanduring carotid occlusion, in 3 it was less andin 1 it was the same. In all of the 3 catswhich showed tachycardia in response to thecombined vascular occlusion after vagotomythe elevation after vagotomy was greater thanthat seen during the carotid occlusion.

These experiments indicated that, while afew cats were carotid sinus-dominant, mostwere pancreatic or viscero-dominant with the

technics of comparison used, since pressor re-sponses of great magnitude occurred in catswith intact carotid sinus or aortic arch re-ceptors, or both, in spite of which a counter-balancing influence from these sites was in-sufficient to inhibit the observed markedelevations of pressure. It appeared, therefore,that the pancreatic and visceral receptorsdominated the reflex regulation of arterialpressure and of heart rate in the majority ofthe cats and could do so for more than the 1.5to 2.0 min. intervals usually studied (fig. 1Bottom).


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o1-FIG. 4. Arterial pressure response to pancreatic arterial occlusion {A), to Trilateral carotid

occlusion (B). Eepeated in the same order in C after 35 nig. tetraethylammonium ion intra-venously. D, E and F show recovery of response to pancreatic arterial occlusion 13, 21 and26 min., respectively, after administration of the drug. Recovery of the response to carotidocclusion (not shown) paralleled the responses seen in D, E. and F. Bilateral cervical vagotomy.Experiment 8.

After obtaining control observations on thepressor responses to carotid and pancreatichypotension in 1 of the 2 cats in which theseresponses were comparable, these observationswere repeated during the maximum effectfollowing the intravenous administration of35 mg. of tetraethylammonium chloride (fig.4). Comparable results were obtained afterthe intravenous administration of 25 mg. ofhexamethoninm chloride.

Although it was thought unlikely that thesimple hydrodynamie effect of the imposedarterial occlusions alone could account forthe observed pressor responses, especiallysince pancreatic blood flow must be small, theexperimental control of this variable wasthought desirable. Figure 5 shows the pressorresponse to pancreatic arterial occlusion bothbefore and after the response to bilateralrenal artery occlusion. Although elevations ofarterial pressure greater than those shown infigure 5C Avere occasionally observed after bi-lateral renal artery occlusion, these were small

relative to those seen after pancreatic or vis-ceral arterial occlusion (fig. 2). There wasonly one response to bilateral renal arteryocclusion which was greater than that shownin figure 2. Figure 5 also shows the reproduci-bility of response obtainable in a steady prep-aration. This type of data suggested the likeli-hood that the observed pressor responses arenot simply a consequence of some type oflocal, nonspecific irritation incident to themechanical arterial occlusion since, if thiswere the case, greater variability would beexpected. This view is further supported bythe results of perfusion experiments describedbelow during which the possibilit}' of a localirritative effect resulting from mechanicalmanipulation was eliminated and those experi-ments in which the' pressor response to vis-ceral hypotension was maintained for longerperiods (fig. 1 Ruttoin).

One experiment was done to examine theresponse to pancreatic vascular occlusion be-fore and after splenectomy. During pan,-


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arterial pressure response to pancreatic arterial occlusion; B, repeat after 6 min.;renal artery occlusion 20 min. after end of B; D, repeat of A. Bilateral cervicalExperiment 7.

creatic vascular hypotension the arterial pres-sure rose from 110/85 to 165/133 before splen-ectomy and from 105/81 to 168/136 afterspleuectomy. An 8 to 12 beat tachycardia oc-curred in both instances. Later in the sameexperiment the entire intestinal tract frompylorus to rectum was rapidly removed (10min.). The pressor response to pancreaticvascular occlusion was 112/87 to 190/153 be-fore enterectomy and 100/80 to 167/137 afterit. Again, a comparable tachycardia occurredin both instances. After subsequent bilateraladrenalectomy 36 min. later, the pressor re-sponse changed from 100/80 to 167/137 andbecame 91/72 to 130/107.

These experiments were aimed at examiningthe reflex effects on arterial pressure of localhypotension in abdominal viscera in generaland in the pancreas in particular. Experi-ments were then designed to examine the re-verse, namely, the reflex effects on arterialpressure of local hypertension in abdominalviscera. This was accomplished by first oc-cluding some or all of the vessels to the areaunder examination which resulted in a re-

flexly induced hypertension as shown above;and then (a) injecting a small volume (1to 2 ml.) of blood forcefully into one of thelocal visceral arteries, (b) withdrawing thesame volume from the femoral artery, and(c) reinjecting it into the visceral artery andthen rapidly repeating this sequence until anew plateau of systemic arterial pressure hadbeen reached.

Although it was possible to accomplish thiswith one of the pancreatic vessels during oc-clusion of the other two and a reflex hypo-tension was noted, the distribution and sizeof the pancreatic vessels made such experi-ments difficult. This type of experiment was,however, readily accomplished during theperfusion technic in the superior mesentericartery with the coeliac and inferior mesen-teric arteries occluded. The results of 3 rep-resentative experiments, of 19 performed in6 cats are shown in figure 6. They indicatethat the elevation of pressure in the vessels.supplied by the superior mesenteric artery isfollowed by systemic hypotension. Other ex-periments were done in which the blood with-


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FIG. 6. A. Preliminary occlusion of coeliac, superior mesenteric and inferior mesentericarteries with elevation of arterial pressure as seen at beginning of tracing. At each signal2 ml. of blood from femoral artery were rapidly injected through the superior mesenterie arterycatheter. Total of 11 2 ml. injections in 30 sec. Bilateral cervical vagotomy. Experiment 17.B. Same experiment. Repeat of A but with 25 1 ml. injections given in 50 sec. C. Sameexperiment but with pressure recorded in a peripheral branch of the superior mesenteric artery.Two milliliters of blood from femoral artery injected at 15 sec. intervals.

drawn from the femoral artery was reiujectedinto the contralateral femoral vein or arteryinstead of the superior mesenteric artery.These experiments resulted in only smallchanges in arterial pressure.

The mesenteric distribution of paciniancorpuscles in the cat is well shown in thestudy of Gammon and Bronk.3 Their presencein the pancreas is shown in figure 7, a photo-micrograph of the pancreas of the cat fromwhich the liemodynamic data was obtainedfor figure 2.*

DISCUSSION

These data lead to the suggestion that thepacinian corpuscles in abdominal visceramediate the reflex circulatory responses oh-

*The authors gratefully acknowledge the assistanceof Dr. George Brecher, Chief, Hemntology Service,National Institutes of Health.

served. This thought is predicated on thecoincidence of (a) the observed circulatoryresponses, (b) the presence in the stimulatedvascular area of receptors (pacinian corpus-cles) known to be sensitive to distortion,4

and (c) the observations of Gammon andBronk3'3 indicating that these receptors emita signal quite as appropriate to the mediationof reflex cardiovascular regulation as are theimpulses arising from the well-studied re-ceptor sites. The possibility, although an un-likely one, must be admitted that Gammonand Bronk might have been recording im-pulses from a receptor other than the paciniancorpuscles that stand out so prominently be-cause of their size. Although the fabric ofproof cannot be woven solely from these coin-cidences, it nevertheless appears reasonableto use this assumption as a working approxi-mation. To do otherwise would require the


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332 SARXOFF, YAMADA

Fio. 7. H & E stain of pancreas of cat number 18(fig. 2). Three paciiiian corpuscles seen in 18 Xmagnification. Insert shows 80 X magnification ofanother corpuscle.

demonstration of the operation in the abdom-inal vascular bed of receptors whose exist-ence is not as yet suspected.

The corpuscles first noted by Vater in 1741°and described in detail by Pacini in 1830were thought by Thoma in 1884, after hefound them in the aortic adventitia, to be re-lated in some way to the regulation of vas-cular tone.7 Schumaker, 25 years later, ob-served that their shape changed with changes inpressure in the mesenteric vascular bed.8 Atabout this time Rainer" and Van de Velde10

and later Collin11 also viewed the function ofthese bodies as being in some way connectedwith vascular tone. Ceelen observed paciniancorpuscles in the pancreas of S9 per cent ofpost-mortem examinations made in man and.interestingly, gained the impression that theyinvolute with age.1- As noted above, Gammonand Bronk*- "' recorded splanchnic action po-tentials synchronous with arterial systole in

the cat and later obtained similar resultsfrom the mesenteric pacinian corpuscles.The}- further observed that, as with the ca-rotid sinus nerve, the over-all discharge fre-quency correlated with the lowering or eleva-tion of arterial pressure after hemorrhage orinfusion. For reasons that are not immedi-ately apparent, they failed to elicit reflexchanges of arterial pressure during variationsof the pressure with which they perfused thesuperior mesenteric artery. Although thereader cannot be entirely clear about theirexperimental conditions, it appears that theydid not simultaneously occlude the coeliac andinferior mesenteric arteries, which have sub-stantial collateralization with the superiormesenteric artery, and thus permitted retro-grade run-off into these vessels rather thanproducing substantial pressure changes in thesmaller vascular distribution of the perfusedsuperior mesenteric artery. They further de-scribed what appear to be preliminary experi-ments with perfusion of the superior mesen-teric artery as a result of which it wassuggested that the afferent impulses to thecentral nervous system arising from themesenteric pacinian corpuscles might serve toregulate visceral blood flow locally and thusadjust the relative distribution of total bloodflow in the organism. Although the experi-ments and data described here are not conso-nant with this suggestion of Gammon andBronk, the parallel which it now appears rea-sonable to draw between the function of thecarotid sinus and that of the abdominal pres-soreceptors relies to a considerable extent onthe neurophysiologic data obtained by them.

Heymans, Bouckaert, Farber and Hsu,13

and Heymans, Bouckaert and Wierzuchow-ski14 studied the possible role of abdominalreceptors in circulatory regulation with a vari-ety of cross-circulation experiments in thedog. A recently published assessment of whatis apparently their present position on thiswork summarizes the matter as follows: "I tis likely that these mesenteric receptors areconcerned with the local distribution of bloodin the abdominal viscera. The reflexes whichthey engender, however, probably contribute


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ill a minor way to the regulation of cardiovas-cular activity which is primarily the concernof the sino-aortic reflexes.'"'1 Other experi-ments in the dog by Erdinan, Ileye and Jung-manu,1"' although aimed primarily at eluci-dating reflexogenic zones in the abdominalaorta proper, yielded interesting and perhapspertinent information. They found that vary-ing the perfusion pressure in a recipient dog"sabdominal aorta receiving blood from a douordog, with or without an intermediate pump,could produce changes in arterial pressure inthe supradiaphragmatic half of the recipientdog even though this half was arterially iso-lated or almost so. Recently Bulekbaeva wasable to elicit reflex effects by varying the per-fusion pressure of the isolated but still inner-vated dog pancreas.10 At this writing a trans-lation of this work is not yet available butfrom the translated abstract it appears tomerit attention:

The reflex effects from the Immorally-isolatedpancreas of dogs under morphine-urethan narcosiswere studied. Pressure stimulation, from changesin the perfusion pressure, resulted in reflexchanges in carotid arterial pressure, in respiratoryfrequency and depth, and in thoracic duct lymphflow. For 50 observations the carotid pressure wasincreased in 28, decreased in 4, variable in 1, andunchanged in 17; the respiration was unchangedin 19, the amplitude increased in 15 and decreasedin 10, the respiratory frequency depressed in 4,increased in 1, and variable in 1; lymph flow in-creased in 41, decreased in 5 and was unchangedin 4. There seemed to be no particular correspond-ence between the various reflx changes. Similarly,chein. stimulation of the pancreas by acetylcholineor insulin produced reflex changes in the carotidpressure, respiration, and lymph flow. Acetylcho-line most often produces an increase in arterialpressure, respiration, and lymph flow. Insulin pro-duced an increase of arterial pressure and oflymph flow, but a decrease of amplitude and fre-quency of respiration. The chemoreceptor reflexchanges of lymph flow seemed independent of thearterial pressure changes.10

The data most relevant to the dog fullyavailable at this time are to be found in theinteresting experiments of Bennati and Ghig-gino.17 They were apparently consistentlyable to produce arterial pressor elevations bythe occlusion of various abdominal arteries,

notably the superior mesenteric artery. Theseresponses were enhanced by vagotomy andblocked by tetraethylammonium chloride. Al-though the pressor responses observed (20mm. Hg) were not as large as those reportedfor the cat in this study, this is compatiblewith the observation that the response to bi-lateral carotid artery occlusion is substantiallymore emphatic in the dog than in the cat.

Since vagotomy did not abolish or diminishthe pressor response to pancreatic or visceralhypotension, the afferent pathway mediatingthe response must, of cou'se, be sympathetic.There is, however, an unfortunately diffuseanatomic distribution of the receptors sus-pected of playing a role in the observed re-flex responses and therefore also the lack of asingle vessel the occlusion of which wouldalone produce it. Further, a definitive attackon the afferent neuronal pathway is made diffi-cult by the absence of a single and separablenerve structure, the cutting of which wouldabolish and the stimulation of which wouldreproduce the physiologic effects under ex-amination. This difficulty is compounded bythe configuration of the sympathetics in thatthe same trunks which contain the afferentfibers mediating this reflex also contain a sub-stantial portion of the efferent fibers whichproduce the response. Such difficulties werenot inherent in the examination of the reflexresponses arising in the carotid sinus and aor-tic arch; this may perhaps at least partiallyaccount for their earlier elucidation.

It is nevertheless possible to draw a reason-ably convincing parallel between the reflexresponses to pressure changes in the carotidsinus and the more diffusely distributed ab-dominal pressoreceptors. With presentlyavailable information this parallel consists of :1. A lowering of pressure in both produces areflex arterial hypertension and tachycardia.2. An elevation of pressure in both producesa reflex arterial hypotension and bradycardia.3. The reflex responses to pressure changes inboth are blocked in parallel by gangl ionicblockade. 4. The action potentials recordedfrom the afferent fibers of both show a cyclicincrease in frequency coincident with arterial


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334 SARXOFF, Y A MAD A

sj-stole and an over-all mean frequency whichis a function of arterial pressure when this ischanged by hemorrhage or infusion.

The observation* that the frequency of dis-charges from the pacinian corpuscles is direc-tionally opposite to that in the carotid sinusnerve when acetylcholine and adrenalin aregiven intravenously do not appear to detractfrom this functional parallel. This does,however, provide ground for the interestingspeculation that, whereas the sinoaortic mech-anism is pre-arteriolar, the receptors underexamination in the above experiments arestimulated at an arteriolar or postarteriolarsite.

With some hesitation, it is suggested thatin the cat the now venerable receptors in thecarotid sinus and aortic arch are dominatedwith regard to their reflex influence on arte-rial pressure by this newly described receptorsystem. No other reasonable interpretationappears consonant with the results observedin the majority of cats in which the pressorresponse and tachycardia was substantiallygreater during either pancreatic or abdominalvisceral vascular hypotension than during bi-latei'al carotid artery occlusion and that intactcarotid sinus and/or aortic arch receptors"permitted" elevations of arterial pressure,during periods of visceral hypotension, whichwere sometimes more than 125 mm. Hg.

SUMMARY

Twenty-four experiments in anesthetizedcats were performed to examine the responseof arterial pressure and heart rate to changingvascular pressures in the blood vessels of theabdominal viscera in general and more par-ticularly in the pancreas. The observationsindicate that receptors in this area, presum-ably the pacinian corpuscles, make a signifi-cant contribution to cardiovascular regulation.

AcKXOWLEDGSIEXT

Grateful acknowledgment is made to Mr. BurtonAlter and Mr. Joseph A. Miles, Jr., for their tech-nical assistance.

SUMMARIO IX INTBRLINGUA

Esseva effectuate vinti-quatro experimentosin gattos anesthesiate con le objectivo de ex-aminar le responsa del tension arterial e delfrequentia cardiac a alterationes del tensionvascular in le vasos sanguinee del visceres ab-dominal in general e del pancreas plus in par-ticular. Le observationes indica que recep-tores in iste area, presumitemente le corpus-culos de Pacini, ha un significative rolo con-tributori in le regulation cardiovascular.

REFERENCES

1. HEYMAXS, C, AXD XEIL, E.: ReflexogenicAreas of the Cardiovascular System. Bos-ton, Little, Brown and Co., 1958.

2. HBRING, H. E.: Die Karotissinus reflexe aufHerz und Gefasse. Leipzig, Steinkopffl,1927.

3. GAMMON, G. D., AXD BROXK, D. W.: The dis-charge of impulses from pacinian corpus-cles in the mesentery and its relation tovascular changes. Am. J. Physiol. 114: 77,1935.

4. HUBBAKD, S. J.: A study of rapid mechanicalevents in a meehanoreceptor. J. Physiol.141: 19S, 1958.

5. GAMMON, G. D., AND BRONK, D. W.: Paciniancorpuscles in mesentery and their relationto vascular system. Proc. Soc. Exper. Biol.& Mod. 31: 788, 1934.

6. SHEEHAN, D.: The clinical significance of thenerve-endings in the mesentery. Lancet 224:409, 1933.

7. THOMA, R.: Ueber die Abhangigkeit der Bin-degewcbsneubildung in der Arterienintimavon den Meehanischen Bedingungen desBlutumlaufes. Virchows Arch. f. Path.Anat. u. Physiol. 95: 294, 1SS4.

S. SCHUMAKER, S.: tJber das Glornus eoccygeumdes Mcnschen und die Glomeruli caudalesder Ssiugetiere. Archiv. f. Mikv. Anat. 71:58, 1907-OS.

9. RAIXER, F. J.: Sur l'existence d'un type geantde corpuscle de Pacini. Comp. rend. Soe. deBiol. 67: 309, 1909.

] 0. VAX DE VELDE, VOX EM. : Die fibrillareStruktur in den Xerveneudorganeu derVogel und der Saugetiere. Anat. Anz. 31:621, 1907.

11. COLLIX, R.: Sur la structure des corpusclesde Vater-Pacini chez le chat. Comp. rend.Soe. de Biol. 85: 511, 1921.

12. CEELEX, W.: L'ber das Vorkommen von Va-ter-Pacini'schen Korperehen am inensch-


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lichen Pankreas und uber eine KrankhafteVerandening derselben. Virchows Arehiv. f.Path. Anat. u. Physiol. 208: 460, 1912.

13. HEYMAXS, C, BOUCKAERT, J. J., FARBER, S.,AND Hsu, F. Y.: Spinal vasomotor reflexesassociated with variations in blood pressure.Am. J. Physiol. 117: 619, 1936.

14. —, BOUCKAERT, J . J. , AND WlEHZUCHOWSKI,M.: Reflexes vasomoteurs medulairres d'ori-gine vasculaLre barosensible. Arch. int.Pharmacol, et Therap. 55: 233, 1937.

15. ERDMANN, W. D., HEYE, D., AND JUNG-

SIANX, H.: Dehuung der Bauchaorta undKreislaufdynaniik. Verhandl. d. Deutsch.ges. £. Kreislaufforschung 22: 113-7, 1956.

16. BULEKBAEVA, L. E.: Reflex effects from re-ceptors of the pancreas on the arterial pres-sure, respiration and lymph flow. Abstracted,Vestnik Akad. Nauk. Kazakh. 13: 10S, 1957.

17. BEXSTATI, D., AND GHIGGKO, C. W.: Efectode hipertension arterial provocado por laoclusioii temporaria de arterias del area es-placnica. Anales de la Facultad de Medi-cina, Montevidea 41: 53, 1956.


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STANLEY J. SARNOFF and SAMUEL I. YAMADAReference to the Pancreas

Evidence for Reflex Control of Arterial Pressure from Abdominal Receptors with Special

Print ISSN: 0009-7330. Online ISSN: 1524-4571 Copyright © 1959 American Heart Association, Inc. All rights reserved.is published by the American Heart Association, 7272 Greenville Avenue, Dallas, TX 75231Circulation Research

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Evidence for Reflex Control of Arterial Pressure from Abdominal