Erythrocytes, or red blood cells, are the most abundant cell type in the equine circulatory system and are responsible for transporting oxygen from the lungs to tissues throughout the body. They also facilitate the return of carbon dioxide from tissues to the lungs for exhalation. Equine erythrocytes are characterized by their biconcave disc shape, which enhances their flexibility and surface area for gas exchange. The lifespan of erythrocytes in horses is approximately 140-150 days, after which they are removed from circulation and broken down, primarily in the spleen. Variations in erythrocyte count, size, and morphology can indicate various health conditions, including anemia, dehydration, and certain metabolic disorders. This page compiles peer-reviewed research studies and scholarly articles that explore the structure, function, and clinical implications of erythrocytes in equine health.
Bowling AT, Wictum E.A fourth allele at the horse erythrocyte phosphohexose isomerase (Phi) locus was proposed to account for phenotypes observed after starch gel electrophoresis and enzymatic staining of red cell lysates from American Saddlebred and Tennessee Walking Horse breeds. The gene was rare, having an estimated frequency of 0.009 in 949 Saddlebreds tested.
Bowling AT, Ewalt-Evans R.Family data are presented for a new allele (Aabdg) in the A system of horse erythrocyte alloantigens which includes factors Aa and Ab traditionally thought to be products of allelic genes. Evidence for incorrect assignment of the codominant factor Ae in the presence of Ab and Ac and the absence of Aa is discussed.
Houle JJ, Hoffmann EM.A passive hemolysis assay system was developed which permitted comparisons of the hemolytic activities of complement (C) from six species. This system employs a single antigen and an antiserum raised in one species. Thus, variations resulting from different target antigens and those inherent in using antibodies (of different affinities and isotypes) raised in a variety of species were minimized. Of the erythrocytes (E) examined, those from horses and guinea pigs were most susceptible to lysis, and either would be suitable, as a tentative choice, for measuring C activity of a previously unstudi...
Harvey JW, Asquith RL, Sussman WA, Kivipelto J.Twenty-one healthy Thoroughbred and Quarter Horse foals were studied from birth until 1 year of age. Foals had access to an iron-supplemented creep feed before weaning and were fed an iron-supplemented concentrate as part of their diet after weaning at 4 months of age. Initial blood samples were taken before foals were allowed to nurse. Serum iron concentration, total iron-binding capacity, and PCV decreased during the foal's first 24 hours of life. Serum iron concentration decreased rapidly from 446 +/- 16 micrograms/dl (mean +/- SE) at birth to 105 +/- 11 micrograms/dl at 3 days of age. Seru...
Valieva TA, Valiev AG, Kulakova SN, Levachev MM.During six weeks rats were given a ration containing, as the only source of fat, lipids of sublimated mare's milk that had been stored with or without antioxidants (quercitrol, bisalcophen) during 3 months. The influence of specific quality of the ration fat on lipids and fatty acid composition of lipids in blood plasma, erythrocytic stroma and liver tissue, as well as on lipid peroxidation in the tissues was studied in the test animals. No changes induced by the lipids from sublimated mare's milk were detected in the parameters studied.
Fincham DA, Mason DK, Paterson JY, Young JD.1. Thoroughbred horses were divisible into five distinct amino acid transport subgroups on the basis of their erythrocyte permeability to L-alanine, measured uptake rates ranging from 5 to 625 mumol l cells-1 h-1 (0.2 mM-extracellular L-alanine, 37 degrees C). 2. Erythrocytes from animals belonging to the lowest L-alanine permeability subgroup (5-15 mumol l cells-1 h-1) (transport-deficient type) exhibited slow nonsaturable transport of this amino acid. In contrast, cells from horses of the four transport-positive subgroups possessed additional high-affinity (apparent L-alanine Km (Michaelis c...
The Journal of heredityMarch 1, 1987
Volume 78, Issue 2 75-80 doi: 10.1093/oxfordjournals.jhered.a110340
Bowling AT, Ryder OA.Ninety-six Przewalski's horses (Equus przewalskii) were blood typed using systems of inherited blood variants known to be highly effective for parentage testing of domestic horses (E. caballus). Sixteen red cell antigenic factors detected using sera prepared by alloimmunization of domestic horses were shown to be inherited in six systems (A, C, D, P, Q, and U) and in the same patterns as domestic horses. Family data confirmed autosomal, codominant inheritance at five loci of serum protein variants (Al, Tf, Xk, Pi, and Es) and three loci of red cell proteins (PGM, PHI, and Hb). One serum protei...
Weiser MG.A multichannel, semiautomated, blood cell counting system (Coulter Counter Model S550) was modified for use in veterinary hematology by increasing both the erythrocyte and leukocyte aperture currents to 225 V and 195 V, respectively, followed by calibration with human blood. It was evaluated by use of 350 samples from dogs, cats, horses, and cows. Values for leukocyte count, erythrocyte count, mean corpuscular volume, and hematocrit generated by the S550 were compared with values generated by an automated multichannel counter with histogram capability and other reference procedures when approp...
Sentsui H, Kono Y.Horse erythrocytes treated with equine infectious anemia virus hemagglutinin were found to be lysed after incubation with fresh horse serum at 37 degrees C. Fresh guinea pig serum induced more efficient hemolysis than horse serum. Direct immunofluorescence test revealed the adsorption of complement factors on the surface of the erythrocytes. Calcium and magnesium ions were necessary for the hemolysis to take place. Antibody against equine infectious anemia virus enhanced the virus-induced complement-mediated hemolysis. These observations indicated that the classical pathway of complement activ...
Moore JN, Mahaffey EA, Zboran M.Heparin was administered subcutaneously 2 times a day for 4 days to 5 horses. An additional group of 5 horses was used as time-matched controls. Significant decreases in PCV, erythrocyte count, and hemoglobin concentration were observed during heparin therapy. The mean corpuscular volume (MCV) of the heparin-treated horses increased to a peak value of 66.1 fl on the last day of treatment. Erythrocyte creatine concentration and glucose 6-phosphate dehydrogenase activity increased moderately during the treatment. These data indicated that the rapid, profound increase in MCV during heparin therap...
Sentsui H, Kono Y.Horse erythrocytes treated with equine infectious anemia virus hemagglutinin were phagocytized by cultivated horse leukocytes (mainly macrophage-like cells and partly polymorphonuclear cells) after incubation with fresh horse serum but not with inactivated horse serum. The phagocytosis began as soon as the erythrocytes were added to the leukocyte cultures, and the majority of the reaction proceeded within 30 minutes. Addition of antiserum showed a slightly suppressing but no enhancing effect on the phagocytosis. Phagocytosis seemed to be caused by the recognition of the third complement compon...
Saltiel A, Gutierrez A, de Buen-Llado N, Sosa C.After parturition, Thoroughbred mares were mated at the first post-partum oestrus (N = 24) or at a subsequent oestrus (N = 12). All mares were examined daily for: oestrous detection, palpation per rectum of the genital tract, vaginoscopic examination and cervico-endometrial cytology. Pregnancy diagnosis was carried out at Days 18, 35 and 45 after mating. An identical first service conception rate of 50% was found in both groups. The number of neutrophils followed a descending profile to only scattered cells at the first post-partum oestrus and in Group II mares remained at this very low level ...
Mahaffey EA, Moore JN.In vitro erythrocyte agglutination developed in 3 hospitalized horses receiving heparin treatment. The agglutination caused artifactual decreases in erythrocyte counts and increases in mean corpuscular volume (MCV) values. Treatment of cell suspensions with trypsin eliminated the agglutination and the changes in erythrocyte count and MCV. Similar abnormalities in erythrocyte counts and MCV have been reported in healthy horses treated with heparin and have been cited as evidence of hemolysis and regenerative anemia.
Radin MJ, Eubank MC, Weiser MG.Anemia was induced in three groups of horses by moderate or severe acute hemorrhage, or by acetyl phenylhydrazine-induced hemolysis (Groups I, II, and III, respectively). Serial hemograms were done on a multichannel automated blood cell counter with histogram capability. Changes in hematocrit, mean cell volume, erythrocyte number, red cell distribution width (RDW), and standard deviation of erythrocyte volume were examined over time. Significant increases in mean cell volume were first detectable by days 17, 20, and 14 and reached maximum by days 43, 41, and 29, in Groups I, II, and III, respe...
Levine RA, Hart AH, Wardlaw SC.Using quantitative buffy coat analysis (QBCA), rapid and accurate measurements can be made of the erythrocyte PCV, total WBC count, and platelet count, and the leukocyte population can be differentiated into total granulocytes (including quantitation of eosinophils), and lymphocytes and monocytes. The QBCA is performed by placing a blood sample (50 to 111 microliters) into a high-precision-bore microhematocrit tube that contains a freely moving, closely fitting, cylindrical plastic float. After centrifugation for 5 minutes, the buffy coat components separate by density. The plastic cylinder fl...
Kuttler KL, Gipson CA, Goff WL, Johnson LW.Nine 4-year-old Arabian geldings were experimentally infected with Babesia equi of European origin. All horses developed detectable parasitemia an average of 30 days after they were inoculated, which was accompanied by a decrease in PCV. The infections were generally mild with no animal deaths. All horses became serologically positive by the indirect fluorescent antibody test within an average of 23 days after they were inoculated and by the complement-fixation test 30 days after they were inoculated.
Allen BV.Results of blood counts have been analysed in three-year-old racehorses in training comprising 77 colt stayers, 27 colt sprinters, 61 filly stayers and 35 filly sprinters. The distributions of haemoglobin, erythrocyte count and haematocrit were significantly higher in colt stayers compared to the other three groups. In fillies these values were also significantly higher in stayers compared to sprinters. The erythrocyte count was significantly higher in filly stayers compared to colt sprinters but there were no significant differences between haemoglobin or haematocrit values in these groups. N...
Amin TM, Sirs JA, Allen BV, Colles CM.A preliminary investigation has been undertaken of blood rheology in horses and ponies, its variation in navicular disease and the changes following treatment with warfarin. Erythrocyte flexibility, measured by a centrifuge packing technique, is higher in horses (30 per cent min-1) than in ponies (23.8 per cent min-1). There are corresponding differences in blood viscosity. The high erythrocyte flexibility in horses is caused by an unknown factor present in plasma. The erythrocyte flexibility in horses with navicular disease is even higher, at 38.5 per cent min-1. Treatment with warfarin reduc...
Contreras A, Martínez R, Devés R, Marusic ET.Marked differences in the activities of three monovalent cation transport systems in horse versus human erythrocytes are reported. Whereas horse erythrocytes exhibit a 6-fold higher sodium-lithium countertransport, the unidirectional flux of potassium through the sodium pump is 3-4 times slower and the sodium-potassium cotransport system cannot be detected. In spite of this, horse and human cells are able to maintain similar Na+ and K+ gradients.
Gordon BJ, Latimer KS, Murray CM, Moore JN.In a continuous-flow centrifugation apheresis technique adapted for blood-component separation and collection in horses, hydroxyethyl starch was not required for erythrocyte sedimentation. The efficacy and separation characteristics of whole blood from 10 horses were evaluated at various gravitational forces (700 to 1,500 rpm), using a constant withdrawal rate (100 ml/min). Maximum leukocyte collection occurred at 700 rpm (P less than 0.01), and optimal neutrophil collection occurred at 700 to 750 rpm (P less than 0.01). Although neutrophil counts decreased and lymphocyte counts remained const...
Yoshihara E, Nakae T.In order to develop the cytotoxic liposome, the cytolytic effect of polycationic liposome was examined. Upon incubation of the stearylamine-containing liposome (stearylamine-liposome) with rabbit erythrocyte, a significant extent of hemolysis was observed. Hemolytic activity of the liposome depends on the amount of stearylamine in the liposome membrane. The plots of the initial rate of hemolysis versus the concentration of stearylamine-liposome showed a sigmoidal curve, suggesting that stearylamine-liposomes act cooperatively on the erythrocyte membrane. Hemolytic activity of stearylamine-lipo...
Gill J, Rastawicka M.In 8 stallions and 6 mares of pure Arabian horses the diurnal changes in the number and certain parameters of red and white blood cells were studied. All of them showed distinct diurnal changes but they did not fulfill the conditions of circadian rhythm according to Halberg. It was found that most of peaks occurred during the dark phase of 24 hr period except the percentage of eosinophils, erythrocyte diameters and cell rate sedimentation. The majority of indices studied showed the higher mean diurnal levels in the mares than in the stallions.
Dinev D, Khubenov KhD.Studied were a total of 85 (33 male and 52 female) donkeys of the Martina Franca breed of the domestic ass, aged from one to twelve years. Stated are the results for the normal values of hemoglobin, hematocrit, ESR, total erythrocyte and leukocyte counts, leukocyte formula, blood sugar, cholesterin, urea, total protein and protein fractions, some micro- and macroelements, and enzymes as tabulated in four diagrams. No differences have been established with regard to sex. It has been found that with young animals the normal values of hemoglobin and erythrocytes are lower than with adult ones. Wi...
Gill J, Kompanowska-Jezierska E.In 34 pure-bred Arabian horses, divided into four groups (Gr. I, 10 pregnant mares; Gr. II, seven barren mares; Gr. III, 10 foals born in 1981; Gr. IV, seven foals born in 1982), seasonal changes in haemoglobin level, haematocrit value, sedimentation rate, red blood cell number and diameter, percentage of erythroblasts and reticulocytes, and index F were studied. Seasonal cyclicity was found in all groups in the haemoglobin level, haematocrit value and RBC diameter. It was also found in the sedimentation rate (PCV) and in index F, but not for the youngest foals (Gr. IV). For the RBC number the...
Easley JR.The erythrogram (erythrocyte histogram) and red cell distribution width (RDW) were evaluated in 5 purebred horses and 1 pony of mixed breeding with experimentally induced anemia. Four horses were studied for 6 weeks after 20% of their estimated blood volume was removed on each of 2 consecutive days (40% total blood loss; acute blood-loss group). Two horses were given acetylphenyl hydrazine IV daily, until acute Heinz body hemolytic anemia was induced; the 2 horses were then evaluated for 6 weeks. One horse and the pony had 20% of their estimated blood volume removed via phlebotomy once each we...
Kuypers FA, Easton EW, van den Hoven R, Wensing T, Roelofsen B, op den Kamp JA, van Deenen LL.The phospholipid composition and the distribution of phospholipids over the two leaflets of the membrane have been investigated for rabbit and horse erythrocyte membranes. Phosphatidylcholine (PC) comprises 39.4% and 41.3% of the total phospholipid complement of the rabbit and horse erythrocytes, respectively. In both membranes the distribution of this phospholipid is asymmetric: 70% of the PC is present in the outer layer of the rabbit membrane and 60% in that of the horse. The major species of this phospholipid class are the (1-palmitoyl-2-oleoyl)- and the (1-palmitoyl-2-linoleoyl)PC. The di...
Engelking LR, Mariner JC.The effect of large-dose heparin therapy on erythrocyte mass depletion in ponies was investigated to determine whether stimulation of reticuloendothelial cell activity and catabolic function would be evidenced by enhanced catabolism of heme to bilirubin. Ponies with chronic external biliary fistula were used to examine biliary excretion of bilirubin both before and after heparin loading (107 U/kg, IV, plus 320 U/kg, subcutaneously) and at maintenance dosages of 320 U/kg given (subcutaneously) at 12 and 24 hours after initial loading with heparin. Results indicated that by 48 hours after ponies...
Tablin F, Weiss L.The equine bone marrow responds to blood loss by increased erythropoiesis, only releasing reticulocytes into the peripheral circulation in severe chronic anemia. We have used morphometric analysis based on electron microscopy of the equine marrow to examine the maturation and release of reticulocytes. Developing red cells in the bone marrows of normal and chronically anemic horses were divided into four stages: early, intermediate, late-stage erythroblasts, and reticulocytes. Morphometric analysis of each stage included volume density of mitochondria per micron3 of cytoplasm, surface area of t...
Fincham DA, Young JD, Mason DK, Collins EA, Snow DH.The amino acid permeability of red blood cells from Equus caballus (thoroughbred, Arab, shire and pony), E przewalskii (Przewalski's horse), E asinus (donkey and mule) and E burchelli (common or plains zebra) was measured. Individual animals exhibited stable but widely differing rates of L-[U-14C]alanine uptake in the range 5 to 1554 mumol (litre cells)-1 h-1 (0.2 mM extracellular L-alanine, 37 degrees C). Of the thoroughbreds tested, 30 per cent had red blood cells which were essentially impermeable to L-alanine (5 to 10 mumol (litre cells)-1 h-1, giving transport rates similar to those found...
Becht JL, Semrad SD.Hematologic parameters change during the first 10 days of life. Erythrocytes increase in number but decrease in size and hemoglobin concentration. The PCV, hemoglobin, and platelet count also decrease. Total blood and plasma volume and, to lesser extent, erythrocyte volume decrease. Normal neonatal foals may have immature neutrophils (up to 5 per cent bands), and their early rapid rise in neutrophil numbers may be accompanied by a lymphopenia. Monocytes, eosinophils, and basophils are all absent or low initially. Infectious processes can cause rapid and variable changes in the leukogram. Howev...
Shull RM.Hemolytic or blood loss anemia was induce in six ponies and red blood cell concentrations of creatine, glucose-6-phosphate dehydrogenase (G-6-PD), lactate dehydrogenase (LDH), and aspartate transaminase (AST) were measured during the ensuing regenerative period. Creatine and G-6-PD levels correlated well and increased concentration of either was good indication of increased erythrogenesis. Erythrocyte LDH levels were of value in assessing the response to hemolytic anemia but not to blood loss anemia. The difference may be, at least in part, the result of differing degrees of regenerative effor...
Smith JE, Kiefer S, Lee M.1. Reduced glutathione regeneration rates as measured in erythro- cytes of ten horses were considerably lower than rates reported in human and sheep erythrocytes.
2. Theactivityofenzymesinvolvedinreducedglutathioneregenerationwas at least three times the observed reduction rate.
3. Despitehighglucose-6-phosphatedehydrogenaseactivity,equineerythro- cytes could not adequately accelerate the hexose monophosphate pathway.
4. Activities of enzymes involved in regulating the glycolytie pathway were less than 5/~moles/min per g hemoglobin; of other glycolytic enzymes greater than 10/~moles/min per...
Medeiros LO, Nürmberger R, Medeiros LF.Erythrocytes from thoroughbred horses were submitted to total (80-90%) and partial (25-40%) oxidation of hemoglobin by sodium nitrite. The ability of these cells to reduce methemoglobin to hemoglobin in the presence of either glucose, glucose plus methylene blue or lactate was investigated. The results were compared with those ones obtained for human erythrocytes. Under total oxidation: the horse erythrocytes need longer incubation time with glucose or glucose plus methylene blue than human erythrocytes for reducing the methemoglobin; methylene blue did not enhance methemoglobin reduction in t...
Vodela JK, Dalvi RR.This study was conducted to determine and compare the activities of glutathione-S-transferase (GST) in red blood cells of cattle, horses, pigs, goats, dogs, rabbits, rats and mice. The highest GST activity was found in mouse red blood cells followed by that of rats, dogs, cattle, pigs, goats and horses with the lowest activity in rabbits. There were significant differences between the GST activities from these various species. The species differences in GST activities correlate with the reported variable responses of the different species to different toxicants since erythrocyte GST plays a si...
Deutsch HF, Bray RP.A study of the erythrocyte carbonic anhydrases of 219 American ponies and 76 riding horses has revealed the presence of five variants of the low-activity CA B isozyme and two variants of the high-activity CA C isozyme. The previously undetected variant of CA C was found only in the pony population and had an allele frequency of 8.9%. A family study of animals possessing the CA B variant A2 showed an unexpected high frequency of inheritance.
Debski B.Erythrocytes, due to their simple metabolic processes, differ greatly from nucleated
cells. Their main metabolite of glycolysis is 2,3-Diphosphoglycerate (2,3-DPG), which
serves also as an energy store for producing ATP in the pyruvate kinase reaction. 2,3-DPG
modifies also the affinity of haemoglobin for oxygen (4). Increase in the concentration of
2,3-DPG, ATP and body temperature, or a decrease in the pH value of blood causes
decreased affinity of haemoglobin for oxygen as blood transverses muscle capillaries,
facilitating oxygen delivery. In equine erythrocytes, the level of ATP is s...
Easley JR.The erythrogram (erythrocyte histogram) and red cell distribution width (RDW) were evaluated in 5 purebred horses and 1 pony of mixed breeding with experimentally induced anemia. Four horses were studied for 6 weeks after 20% of their estimated blood volume was removed on each of 2 consecutive days (40% total blood loss; acute blood-loss group). Two horses were given acetylphenyl hydrazine IV daily, until acute Heinz body hemolytic anemia was induced; the 2 horses were then evaluated for 6 weeks. One horse and the pony had 20% of their estimated blood volume removed via phlebotomy once each we...
Gingerich DA, Mia AS.Thirty adult horses were used to compare the toxicity and cholinesterase (ChE) inhibition of various dosages of a combination anthelmintic, trichlorfon + mebendazole. Single oral doses of up to 5 times the effective dosage (39.7 mg of trichlorfon and 8.8 mg of mebendazole/kg of body weight) did not result in deaths. Horses given a placebo and horses treated at the recommended dosage rate showed few or no side effects, whereas horses given higher dosages showed dosage-related increases in the severity of clinical signs of organophosphate toxicosis. Dosage-related inhibition of erythrocyte ChE a...
Bowling AT, Wictum E.A fourth allele at the horse erythrocyte phosphohexose isomerase (Phi) locus was proposed to account for phenotypes observed after starch gel electrophoresis and enzymatic staining of red cell lysates from American Saddlebred and Tennessee Walking Horse breeds. The gene was rare, having an estimated frequency of 0.009 in 949 Saddlebreds tested.
INGRAM DG.A method is described for the quantitative measurement of the reactions between sensitized cells and horse complement and between alexinated cells and conglutinin. The method is laborious but its application has allowed the determination of the optimal times of the reactions at various temperatures. The results obtained in these experiments indicate that the alexinated configuration with which conglutinin and immuno-conglutinin react is not one of the recognized intermediates formed during the process of immune haemolysis.
Murayama JI, Takeshita K, Tomita M, Hamada A.Crude glycophorin fraction was prepared from horse erythrocyte membranes by extraction with lithium diiodosalicylate and partition in aqueous phenol. Two glycophorins, designated glycophorins HA and HB, were isolated by two different techniques: preparative gel electrophoresis in the presence of sodium dodecyl sulfate and ion-exchange chromatography in the presence of the nonionic detergent Ammonyx LO. Each glycophorin formed at least two bands on gel electrophoresis, which corresponded to a dimeric form and a monomeric form. Glycophorin HA, the major component, had a blocked amino-terminus an...
Knych HK, Wilson WD, Vale A, Kass PH, Arthur RM, Jones JH.Due to the high prevalence of EIPH in racehorses and its potential impact on the horse's health, furosemide administration is permitted up to 4-h prior to post time in most North American racing jurisdictions. Anecdotal reports suggest that administration of furosemide 24-h prior to strenuous exercise may be equally effective in decreasing the severity of EIPH. Objective: To 1) compare the efficacy of furosemide in reducing the presence and severity of EIPH when administered 4- or 24-h prior to strenuous exercise 2) characterise electrolyte and blood parameters following administration of furo...
Jarvis SM, Harris RC.The inward transport of two purines, adenosine and hypoxanthine, at 37 degrees C by horse erythrocytes was compared. No mediated transport of adenosine was detected in horse erythrocytes, nor was saturable, high-affinity binding of the potent facilitated-diffusion inhibitor nitrobenzylthioinosine demonstrable in horse erythrocyte membranes. In contrast, erythrocytes from most horses possessed a saturable sodium-dependent hypoxanthine transporter (apparent K(m), 100 +/- 28 microM; Vmax, 0.20 +/- 0.08 mmol (l cells)-1 h-1; means +/- S.E.M., n = 5). Guanine inhibited hypoxanthine influx (apparent...
Lazary S, Gerber H, de Weck AL, Arnold P.A new, non-MHC linked alloantigenic membrane antigen on the equine lymphocytes is described. This antigen was characterized with alloantisera in the two-stage microcytotoxicity test and designated as ELy-1 antigen. The frequency of ELy-1 antigen positive animals in various populations is close to 50%. ELy-1 shows an autosomal, dominant inheritance. Since an allelic antigen (s) could not be demonstrated in family studies, it is assumed that only two alleles ELy-1+ and ELy-1- exist. The ELy-1 antigen in positive animals is expressed on both T and B lymphocytes but it is not present on erythrocyt...
Funkquist P, Wagner PD, Hedenstierna G, Persson SG, Nyman G.In order to evaluate the pulmonary gas exchange during exercise in Standardbred trotters with red cell hypervolaemia (RCHV), 12 horses with RCHV were compared with 9 normovolaemic (NV) horses. VO2 and VCO2 were determined with an open bias flow system. Cardiovascular and haemodynamic data were recorded during exercise at 4 different speeds on a treadmill. Pulmonary gas exchange was assessed by conventional blood gas variables (arterial and mixed venous blood gas tensions), and the ventilation-perfusion distribution VA/Q was estimated by the multiple inert gas elimination technique. VA and AaDO...