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Peripheral Circulation

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Abstract

Blood flow (BF) increases with increasing exercise intensity in skeletal, respiratory, and cardiac muscle. In humans during maximal exercise intensities, 85% to 90% of total cardiac output is distributed to skeletal and cardiac muscle. During exercise BF increases modestly and heterogeneously to brain and decreases in gastrointestinal, reproductive, and renal tissues and shows little to no change in skin. If the duration of exercise is sufficient to increase body/core temperature, skin BF is also increased in humans. Because blood pressure changes little during exercise, changes in distribution of BF with incremental exercise result from changes in vascular conductance. These changes in distribution of BF throughout the body contribute to decreases in mixed venous oxygen content, serve to supply adequate oxygen to the active skeletal muscles, and support metabolism of other tissues while maintaining homeostasis. This review discusses the response of the peripheral circulation of humans to acute and chronic dynamic exercise and mechanisms responsible for these responses. This is accomplished in the context of leading the reader on a tour through the peripheral circulation during dynamic exercise. During this tour, we consider what is known about how each vascular bed controls BF during exercise and how these control mechanisms are modified by chronic physical activity/exercise training. The tour ends by comparing responses of the systemic circulation to those of the pulmonary circulation relative to the effects of exercise on the regional distribution of BF and mechanisms responsible for control of resistance/conductance in the systemic and pulmonary circulations. © 2012 American Physiological Society. Compr Physiol 2:321‐447, 2012.

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Figure 1. Figure 1.

Central hemodynamics during incremental exercise to exhaustion. Left column: cardiac output, heart rate (HR), stroke volume, mean arterial (•) and central venous (▾) pressures, and systemic vascular conductance. Right column: arterial O2 content, systemic O2 delivery, systemic a‐vO2 difference, systemic O2 extraction, and systemic O2 uptake during incremental exercise to exhaustion plotted against the relative increase in power output. Data are means ± SEM for eight subjects. * Lower than 80% of peak power, P < 0.05. Adapted from with permission of the Physiological Society and Wiley‐Blackwell.

Figure 2. Figure 2.

Regional distribution of cardiac output during various intensities of exercise expressed as % maximal oxygen consumption. Bar graphs on the left illustrate total cardiac output (sum of all tissue flows) and the regional distribution of cardiac output to all tissues with increasing exercise intensity. Bar graphs on the right show an expansion of regional blood flow (BF) data to all tissues except skeletal muscle tissue, to enhance appreciation of the changes in BF to the other tissue with exercise (Note the change in the BF axis for the graphs on the right). Data are estimated for human values using Rowell's text book (also estimates for humans) and our data for regional BF in pigs .

Figure 3. Figure 3.

Schematic drawing of an arteriole (top) and of endothelium, VSM and parenchymal cell (at the bottom) illustrating mechanisms for control of vasomotor tone and diameter. Adapted from Figure of with permission of the American Physiological Society. Neurohumoral, endothelial, and metabolic influences are detailed in the bottom part of the figure. Abbreviations: KCa, calcium‐activated K+ channel; KATP, ATP‐sensitive K+ channel; Kv, voltage‐gated K+ channel; KIR, inward rectifying K+ channel; Trp, transient receptor potential channels; O2, oxygen; ATP, adenosine triphosphate; NO, nitric oxide; TXA2, thromboxane A2 and receptor; 5HT, 5‐hydroxytryptamine and receptor; P2, purinergic type 2 receptor; M, muscarinic receptor; H1 and H2, histamine type 1 and 2 receptors; B2, bradykinin type 2 receptor; ECE, endothelin‐converting enzyme; bET‐1, big endothelin‐1; ET‐1, endothelin‐1; eNOS, endothelial nitric oxide synthase; L‐arg, L‐arginine; COX‐1, cyclooxygenase 1; CYP450, cytochrome P450; ACE, angiotensin‐converting enzyme; AI, angiotensin I; AII; angiotensin II; AT1, angiotensin type 1 receptor; AT2, angiotensin type 2 receptor; ETA, endothelin type A receptor; ETB, endothelin type B receptor; PG, prostaglandins; AA, arachidonic acid; EDHF, endothelium‐derived hyperpolarizing factor; O2, superoxide anion; VGCC, voltage‐gated calcium channels; IP, prostacyclin receptor; EETs, epoxyeicosatrienoic acids; HETEs, hydroxyeicosatetraenoic acids; H2O2, hydrogen peroxide; α1, α1‐adrenergic receptor; α2, α2‐adrenergic receptor; β2, β2‐adrenergic receptor; ACh, acetylcholine; NE, norepinephrine; NPY, neuropeptide Y; P1, purinergic type 1 receptor; and Y1, neuropeptide Y receptor.

Figure 4. Figure 4.

Segmental distribution of control mechanisms throughout the arterial microcirculation, redrawn from with permission of the American Physiological Society.

Figure 5. Figure 5.

Relationships between cardiac output (solid symbols) and two‐legged blood flow (open symbols), systemic vascular conductance (solid symbols) and two‐legged vascular conductance (open symbols), and systemic O2 delivery (solid symbols) and two‐legged O2 delivery (open symbols) versus the increases in systemic oxygen consumption during incremental exercise to exhaustion. Data are means ± SEM for six subjects. Adapted from with permission of the Physiological Society and Wiley‐Blackwell.

Figure 6. Figure 6.

Vasodilation of arterioles and increases in blood flow (BF) can be detected within 1 s after contraction and BF increases with a similar time course in humans. (A) A record of arteriolar diameter and tissue pO2 during single fiber stimulation showing rapid dilation and decrease in pO2. Adapted from with permission of the American Physiological Society. (B) Muscle BF response to a mild‐intensity dynamic exercise bout as measured in dogs with ultrasonic flow probes. Note the immediate increase in BF at the inititation of dynamic exercise (at the arrow). Adapted from with permission of the American Physiological Society. (C) Representative diameter records in 2A branch arterioles after single contractions for durations of 400 and 600 ms of hamster retractor muscle. Adapted from with permission of the American Physiological Society. (D) Arteriolar dilation following a 250 ms tetanic contraction of 3 to 5 fibers of hamster cremaster muscle. Graphed from data from . (E) Femoral artery BF in transition from rest to passive exercise and to voluntary, one‐legged, dynamic knee‐extensor exercise at 10 W (n = , 30 W (n = , 50 W (n = , and 70 W (n = . Voluntary exercise started at 0 time. BF was significantly greater than resting BF during passive exercise and during all time points during voluntary exercise and from passive to voluntary exercise. Adapted from Rådegran and Saltin with permission of the American Physiological Society. (F) Femoral artery BF velocity (Vmean) in transition from rest to passive exercise and to voluntary, one‐legged, dynamic knee‐extensor exercise at 50 W. Knee‐extensor force (F), intramuscular pressure (IMP), arterial (BPa) and venous (BPv) blood pressure and HR in beats per minute (BPM) are presented. Onset of voluntary exercise is indicated by an arrow on the tracing for all parameters in panel F. Adapted from Rådegran and Saltin with permission of the American Physiological Society.

Figure 7. Figure 7.

Results of immunoblot analysis of eNOS expression throughout arteriolar trees (each branch order) of gastrocnemius and soleus muscles of rats chronically endurance trained (ET; panels A,C,D) or chronically trained with an interval sprint training program (IST; panel B). Panels A and B present results for the gastrocnemius muscle and panel C presents eNOS expression for the soleus muscle. Panel D presents results for expression of superoxide dismutase (SOD) in feed arteries and arterioles of the soleus muscle. To clarify effects of training, results are expressed relative to protein content of arteries from sedentary rats (SED) as ET/SED or IST/SED. GFA = gastrocnemius feed artery, the first branch from the feed artery is the 1A arteriole and the branches of the 1A are subsequent 2A branches etc. WG = white portion of the gastrocnemius muscle and RG = the red portion of the gastrocnemius muscle. SFA is the soleus feed artery and the branches of the arterioles in the soleus are S1A, S2A, etc. Note that ET resulted in significant increases in eNOS protein in the arterioles feeding red muscle while IST training exhibited increased eNOS in the GFA, and WG4A and RG5A arterioles. Endurance training did not alter eNOS or SOD‐1 expression in the soleus arterioles. Endurance trained results are from McAllister et al. and those for the IST rats are from Laughlin et al. . * = value different from 1.0 with p < 0.05; + = value different from 1.0 with p < 0.1.

Figure 8. Figure 8.

Distribution of total CO among legs, respiratory muscles (RM), and other metabolically active tissues (skin, heart, brain, kidneys, and liver) during exercise at o2 max. Adapted from with permission of the American Physiological Society.

Figure 9. Figure 9.

Bar graphs of calf blood flow at rest and during maximal hyperemia (post ischemic exercise) in sedentary and trained younger and older women (top panel) and men (bottom panel). Maximal hyperemia was elicited by ischemic exercise to fatigue and measuring flow after release of the occlusion. Data are mean ±SD. *P < 0.01, **P < 0.05 versus younger subjects of corresponding gender and training status. †P < 0. 01, ‡P < 0.05 versus sedentary subjects of corresponding age and gender

. § P < 0.01, §§ P < 0.05 versus men of similar age and training status. Data taken, with permission, from Martin et al. .

Figure 10. Figure 10.

Arm blood flow (ABF) and a‐vO2 difference during arm cranking to exhaustion in rowers and average, fit subjects. A + L, addition of leg exercise to arm cranking in the rowers. Values are means ± SE for eight average fit subjects and seven rowers. Adapted from Volianitis et al. with permission of the American Physiological Society.

Figure 11. Figure 11.

Upper limb (arm) and lower limb (leg) muscle blood flow (BF) (top) and vascular conductance (bottom) from rest to peak exercise. Submaximal (SM) exercise was expressed in terms of relative exercise intensity, SM1, SM2, and SM3 (approx. 40%, 60%, and 80%, of peak power respectively). Values are mean ± SEM. *; significantly different from controls. §; significantly different from swimmers. Irrespective of the group, BF max and conductance was increased significantly from rest to peak exercise. Adapted from Walther et al. with permission of Acta Physiologica.

Figure 12. Figure 12.

Changes in arteriolar density do not explain changes in blood flow (BF) capacity following interval sprint training (IST) and endurance exercise training (ET). Results illustrate the effects of training on oxidative capacity (cytochrome c concentrations), BF capacity, capillary density, and arteriolar density in rat skeletal muscles. All data are expressed as a % increase above respective SED values. The IST training consisted of 10 weeks of 6 training bouts/day, 5 days/week, with each rat running 60 m/min up a 15% incline for 2.5 min with 4.5 min of rest between bouts. The ET training program consisted of 10 to 12 weeks of treadmill running at 30 m/min, 60 min/day, 5 days/week. The top row of data are for the white portion of the gastrocnemius muscle (GW), the middle row for the red portion of the gastrocnemius muscle (GR), and the bottom row of data are for the soleus muscle. These data indicate that IST increased oxidative capacity and BF capacity most in the GW muscle and these changes were correlated with increases in capillary density and small increases in arteriolar density. In contrast, ET increased oxidative capacity and BF capacity most in the soleus and GR muscle but these changes were not correlated with changes in capillary or arteriolar density in the soleus muscle. In the GR muscle, ET increases in BF capacity were similar to increases in arteriolar density. Of interest, IST increased oxidative capacity and BF capacity in the GR muscle but neither capillary nor arteriolar density were altered in this muscle tissue by IST. These data were taken from .

Figure 13. Figure 13.

Leg blood flow responses to graded upright cycle ergometer in younger and older healthy adults. Most results indicate that the hyperemic response to large muscle dynamic exercise is blunted in older humans. Adapted from Proctor and Parker with permission of the Microcirculatory Society.

Figure 14. Figure 14.

Myocardial oxygen balance in awake dogs at rest and during four incremental levels of treadmill exercise. The increase in myocardial oxygen consumption was principally accounted for by an increase in coronary blood flow with only modest contributions of increase in hematocrit and oxygen extraction. MVO2, myocardial oxygen consumption, Hct, hematocrit, ArtSO2, arterial oxygen saturation, CVSO2, coronary venous oxygen saturation. Data are from von Restorff et al. , and have been presented as mean ± SE. *P < 0.05 versus rest. Modified from with permission of the American Physiological Society.

Figure 15. Figure 15.

Relations between HR and left ventricular myocardial blood flow (LVMBF) at rest and during treadmill exercise in dogs 1258, 1259, 1391, 1392, , swine and horses . Data from humans were obtained principally from young healthy male subjects performing upright bicycle exercise 1492, 1493). Data from rats have been added (solid circles) to illustrate that the high LVMBF values in this species are the result of the high heart rates, so that the rat data fall close to the regression line for the human data. Modified from with permission of the American Physiological Society.

Figure 16. Figure 16.

Relation between M o2 and coronary venous oxygen tension (left panel) and oxygen saturation (right panel) at rest and during treadmill exercise in humans , swine , horses , and dogs . Note that exercise does not alter coronary venous oxygen tension in swine, whereas it is already reduced at low levels of exercise in dogs. Humans and horses demonstrate an intermediate oxygen tension response. During exercise oxygen saturation decreases in horses and humans (similar to dogs) whereas in swine saturation is not affected, suggesting a rightward shift of the hemoglobin‐oxygen dissociation curve during exercise in humans and horses. Also note that despite similar resting coronary venous oxygen tensions, horses and humans have higher hemoglobin‐oxygen saturations as compared to swine and dogs, consistent with the higher P50 values reported in the latter species . Data are mean ± SE. *P < 0.05 versus corresponding rest. See text for further explanation. Modified from with permission of the American Physiological Society.

Figure 17. Figure 17.

Coronary blood flow and hemodynamic responses to treadmill exercise in dogs. Modified from Khouri et al.. L. Circ. = left circumflex coronary artery; Cor. = coronary; Syst. = systolic; Diast. = diastolic. See text for further explanation. Modified from with permission of the American Physiological Society.

Figure 18. Figure 18.

Coronary pressure‐flow relation in the dog heart under conditions of maximal coronary vasodilation with intracoronary adenosine (50 μg/kg/min). Shown are the relations at rest and during three incremental levels of treadmill exercise. Note the rightward shift of the pressure‐flow relation with an increase in the zero flow pressure (Pzf) intercept. See text for further explanation. Data are from Duncker et al. and have been presented as mean ± SE. *P < 0.05 versus corresponding rest. Modified from with permission of the American Physiological Society.

Figure 19. Figure 19.

Graph showing a schematic drawing of the intramyocardial microvasculature (upper panel) and the extravascular forces acting on the coronary microvasculature during diastole (left lower panel) and systole (lower right panel). Abbreviations: PIM = intramyocardial pressure; PLUMEN = pressure in left ventricular lumen; PPERI = pressure in pericardial space; PPERI = pressure in pericardial space; Left ventricular lumen pressure. See text for further explanation. Modified from with permission of the American Physiological Society.

Figure 20. Figure 20.

Distribution of left ventricular myocardial blood flow in the dog at rest (open symbols) and during exercise (solid symbols) in the presence of intact vasomotor tone (circles) and during maximum vasodilation with intracoronary adenosine (squares). Data are from Duncker et al. and have been presented as mean ± SE. *P < 0.05 versus corresponding rest. Dot inside symbol denotes significant (P < 0.05) increase in flow produced by adenosine. Modified from with permission of the American Physiological Society.

Figure 21. Figure 21.

Relation between heart rate (HR) and left ventricular subendocardial to subepicardial blood flow (BF) ratio (LV ENDO/EPI) at rest and during treadmill exercise in dogs , swine 1004, 1005, 1205, 1206) and horses , measured with microspheres 15 μm in diameter. No data on transmural distribution of left ventricular BF are available in humans. Note that exercise results in a modest decrease in ENDO/EPI ratio, but this typically does not decrease below 1.0 even during heavy exercise. Also note that ENDO/EPI ratios are lower and decrease more during exercise in horses (LV E/E = −1.37•HR + 1.30) than in either swine (LV E/E = −1.17•HR + 1.38) or dogs (LV E/E = −1.06•HR + 1.41). Modified from with permission of the American Physiological Society.

Figure 22. Figure 22.

Relation beween myocardial oxygen consumption (MVO2) and coronary venous oxygen tension (CVPO2) in the right ventricle (RV) and the left ventricle (LV) in dogs during treadmill exercise. Note the lower levels of MVO2 and higher levels of CVPO2 in the RV compared to the LV. Data are from Hart et al. for RV data and from Gorman et al. for LV data. Data have been presented as mean ± SE. See text for further explanation. Modified from with permission of the American Physiological Society.

Figure 23. Figure 23.

Left panels show the effect of mixed α1‐/α2‐adrenergic receptor blockade with phentolamine on the relation between myocardial oxygen consumption (MVO2) and coronary venous oxygen tension (CVPO2) in the left ventricles of dogs and swine during treadmill exercise. Right panels show the effect of combined α1‐/α2 and β1‐/β2‐adrenergic receptor blockade with phentolamine and propranolol in swine and dogs . Data are mean ± SE. *P < 0.05 versus corresponding control or versuscorresponding phentolamine alone. See text for further explanation. Modified from with permission of the American Physiological Society.

Figure 24. Figure 24.

Integrative metabolic control of coronary vasomotor tone in dogs (left panels) and swine (right panels) at rest and during treadmill exercise. Shown are the effects of adenosine receptor blockade with 8‐phenyltheophylline [8PT 5 mg/kg iv ; upper panels], the effects of KATP channel blockade with glibenclamide [Glib; 50 μg/kg/min ic , or 3 mg/kg iv ] and additional adenosine receptor blockade (middle panels), and the effects of NO synthase inhibition with N‐nitro‐L‐arginine [LNA, 1.5 mg/kg ic or 20 mg/kg iv ] and additional adenosine receptor blockade and KATP channel blockade (lower panels) on the relation between myocardial oxygen consumption (M o2) and coronary venous oxygen tension (CVPO2) in the left ventricles. Data in dogs are from references . Swine data are from Merkus et al. . Data are mean ± SEM. *P < 0.05 effect of 8PT; P < 0.05 effect of glibenclamide; P < 0.05 effect of LNA. See text for further explanation. Modified from with permission of the American Physiological Society.

Figure 25. Figure 25.

Effects of exercise training in swine on DNA labeling in capillaries (top left panel), sprouting of new capillaries (top middle panel), % labeling of sprouts (top right panel), capillary diameter (bottom left panel), capillary density (bottom middle panel), and coronary transport reserve (CTR; bottom right panel). Data are from White et al. and have been presented as mean ± SE. *P < 0.05 versus 0 week time point (sedentary swine). See text for further explanation. Modified from with permission of the American Physiological Society.

Figure 26. Figure 26.

Graph summarizing the structural and functional coronary microcirculatory adaptations to chronic exercise training. ACh = acetylcholine; M = muscarinic receptor; NE = norepinephrine; α1 = α1‐adrenergic receptor. β2 = β2‐adrenergic receptor. Modified from with permission of the American Physiological Society.

Figure 27. Figure 27.

Indices of cerebral blood flow (BF) and oxygenation during progressive exercise expressed as % of baseline value. Arterial carbon dioxide tension (Paco2). Frontal lobe oxygenation (Sco2) determined with near‐infrared spectroscopy. The calculated average cerebral mitochondrial oxygen tension (PmitoO2). The cerebral oxygen consumption rate (CMRO2). Middle cerebral artery mean flow velocity (MCA Vmean). Cerebral BF expressed as grey matter flow (F1) and as the average flow (ISI) during cycling exercise. ISI and F1 were not determined during maximal exercise. Data from .

Courtesy of T. Seifert, the Copenhagen Muscle Center, University of Copenhagen.
Figure 28. Figure 28.

Linear relationships between middle cerebral artery mean flow velocity (MCA Vmean) and cardiac output ( ) at rest and during exercise. • Rest ○ Exercise. Data are means (SEM). Modified from with permission of the Physiological Society and Wiley‐Blackwell.

Figure 29. Figure 29.

Effects of leg muscle tensing on arterial pressure oscillations and low‐frequency power spectrum. Left: original recording. Middle: beat‐to‐beat values. Right: fast Fourier power spectrum of mean arterial pressure. The amplitude of the oscillations in arterial pressure during standing (upper panel) is attenuated during leg tensing (lower panel) with a reduction in the low‐frequency (0.07 to 0.15 Hz) power spectrum. Modified from, with permission, the Ph.D. Thesis of Pott, 2003 (1096, 1097).

Figure 30. Figure 30.

Regional cerebral blood flow (rCBF) by 133Xe SPECT at rest and during static handgrip exercise before and after regional blockade in 1 subject. Scale (right) represents absolute flow values. There is an increase in flow to the motor sensory area in control contraction condition (upper right) compared with rest or with contractions after axillary blockade. Premotor rCBF increased from 55 to 60 ml/100 g/liter/min (n = and motor sensor rCBF from 57 to 63 ml/liter/100 g/liter/min to both the ipsilateral and contralateral sides during handgrip exercise before, but not after, axillary blockade. Modified from Friedman et al. with permission of the American Physiological Society.

Figure 31. Figure 31.

Schematic description of the thermoregulatory control of skin blood flow (BF) as modified by moderately intense exercise. The relation of skin BF to internal temperature is affected, relative to resting conditions, in at least three ways by exercise: a vasoconstrictor response at the onset of dynamic exercise (A), an increase in the internal temperature threshold at which skin BF begins to increase (B), and a leveling off, or plateau, in skin BF above an internal temperature of 38°C at a level well below maximal (C). Exercise exerts these effects through the vasoconstrictor system for the initial vasoconstriction and through inhibiting the active vasodilator system for the increased threshold and for the plateau. At rest, the plateau only occurs as skin BF approaches maximal vasodilation. Modified from Gonzalez‐Alonso et al. 2008 with permission of the Physiological Society and Wiley‐Blackwell.

Figure 32. Figure 32.

Heterogeneity of blood flow (BF) to various bone tissue in anesthetized dogs. Values below the bars reflect the number of observations for the mean value illustrated. Epiphysis is from the femur inferior epiphysis; “diaphysis represents 28 samples of femur diaphysis and 8 samples of humerus diaphysis.” BFs from the two diaphyseal regions were not significantly different (P > 0.05; “red marrow” is from the femur medulla, and “yellow marrow” represents samples from the medullas of the tibia and mandible. Analysis of variance and Tukey's test indicate the following significant differences: sternum, rib, ilium, and epiphysis > diaphysis and yellow marrow; sternum > rib; red and yellow marrow > diaphysis; red marrow > yellow marrow. Modified from Gross et al. 1979 with permission of the American Physiological Society.

Figure 33. Figure 33.

Relationship between splanchnic blood flow (BF) and exercise intensity (expressed as percent o2 max) at normal temperature (25.6°C, solid line) and during heat stress (43.3°C, dashed line). The reduction of splanchnic BF during exercise is proportional to the relative exercise intensity (% o2 max) and is further reduced by environmental heat stress resulting in ∼20% greater reductions in splanchnic BF at a given relative exercise intensity. Modified from Rowell, 1993 which was adapted from Rowell 1965 with permission of the American Physiological Society.

Figure 34. Figure 34.

Exercise‐induced reductions in both splanchnic blood flow (BF) and splanchnic blood volume are nonuniformly distributed across the splanchnic organs. (A) Nonuniform changes in splanchnic BF during and 2 minutes after termination of cycling. The decreases in total splanchnic BF (SBF) (top panel) during moderate cycling exercise involve greater reductions in celiac artery BF (dotted line; middle panel) than in the superior mesenteric artery BF (solid line; middle panel) owing to differential increases in artery resistance (R) in the celiac (dotted line) and superior mesenteric artery (solid line) vascular beds (bottom panel). For both BF and R data in the middle and bottom panels, respectively; continuous line = superior mesenteric artery and the dashed line = the celiac artery. * = different from the preceding value, P < 0.05; # = different from rest, P < 0.05. Data from Perko et al. with permission of the American Physiological Society. (B) Nonuniform changes in splanchnic blood volume. Splanchnic blood volume redistribution during zero‐load (ZLC) and graded cycling exercise (50%‐100% maximal exercise) and 5 min of recovery from exercise. Note that the volume redistribution involves a greater reduction of splenic than hepatic and renal blood volume and that bowel blood volume did not significantly change from baseline values during or following exercise. Values are % basal volume, mean ± SE. Data taken from Table of Flamm et al. 1990 .

Figure 35. Figure 35.

Exercise training alters the redistribution of blood flow (BF) to kidney and splanchnic tissues during exercise. Top panel: baseline/resting BF values represent sedentary rats standing quietly on the treadmill. Middle panel: exercise BF values for sedentary rats at 15 min of treadmill running 30 m/min. Bottom panel: exercise BF values for exercise trained rats at 15 min of treadmill running 30 m/min. Baseline BFs were similar between sedentary and trained rats. As shown in the middle panel, exercise caused BF reductions to all organs, except the liver in sedentary animals. In contrast, results presented in the bottom panel indicate that trained animals exhibit an attenuated reduction of splanchnic and renal BFs during acute exercise. Values are mean ± SE. Liver BF represents hepatic artery BF only. Data are from Armstrong and Laughlin and McAllister .

Figure 36. Figure 36.

Graded cycling exercise markedly increases renal vascular resistance thereby reducing renal blood flow (inferred from reduced renal venous outflow). Workloads were 30%, 60%, and 80% to 90% o2 max. Values are mean ± SE. The *, **, and *** indicate values significantly different from rest, with p < 0.05, 0.01, and 0.001, respectively. Adapted from Tidgren et al. with permission of the American Physiological Society

.

Figure 37. Figure 37.

Changes in pulmonary artery pressure (PAP, open circles in left panels), left atrial pressure (solid circles in left panels) and pulmonary vascular resistance (PVR, right panels) as a function of estimated exercise intensity in dogs , swine , sheep , humans , and horses . Note that PVR is inversely related to body weight (dogs ∼15 kg; swine and sheep ∼25 kg; humans ∼75 kg; and horses ∼400 kg). Exercise intensity is calculated based on heart rate (dogs, sheep, and horses) and/or body oxygen consumption (swine and humans). From Merkus et al. with permission of Elsevier.

Figure 38. Figure 38.

Integrated endothelial control of pulmonary vascular tone in swine (554). The vasodilator effect of ETA/ETB receptor blockade with tezosentan was enhanced by prior inhibition of eNOS (middle panels), but not by COX inhibition (bottom panels) at rest and during exercise in swine. These findings indicate that NO limits the influence of ET on the pulmonary vasculature. B o2: body oxygen consumption; PAP: pulmonary artery pressure; PVR: pulmonary vascular resistance. *P < 0.05 versus control, †P < 0.05 effect of tezosentan different after eNOS/COX inhibition. Adapted from Merkus et al. with permission of Elsevier.

Figure 39. Figure 39.

Autonomic control of pulmonary vascular tone in swine . α‐Adrenoceptor blockade results in vasodilation (top), whereas β‐adrenoceptor blockade results in vasoconstriction at rest and during exercise (middle). Hence, the sympathetic nervous system exerts no net effect on pulmonary vascular tone. M‐receptor blockade results in pulmonary vasodilation at rest that wanes with increasing exercise intensity (bottom), suggesting that withdrawal of the vasoconstrictor influence exerted by the parasympathetic nervous system contributes to exercise‐induced pulmonary vasodilation. B o2: body oxygen consumption; PAP: pulmonary artery pressure; PVR: pulmonary vascular resistance. *P < 0.05 versus control. From Merkus et al. with permission of Elsevier.

Figure 40. Figure 40.

Schematic regulation of the pulmonary vascular tone at rest and during exercise in the healthy pulmonary circulation. NO: nitric oxide; PGI2: prostacyclin; βAR: β adrenoceptor; MR: muscarinic receptor; Ado: adenosine; ANP: atrial natriuretic peptide; KATP, KCa2+ and KV: ATP, Ca2+ and voltage‐sensitive K‐channels; ETA: endothelin A receptor; ETB: endothelin B receptor; αAR: α‐adrenoceptor; Ang‐II: angiotensin II; TXA2: thromboxane A2; 5‐HT: 5‐hydroxytryptamine; PDE5: phosphodiesterase 5; ROS: reactive oxygen species; +: significant influence; −: no influence; ?: probable influence (not yet proven). See text for further explanations. From Merkus et al. with permission of Elsevier.



Figure 1.

Central hemodynamics during incremental exercise to exhaustion. Left column: cardiac output, heart rate (HR), stroke volume, mean arterial (•) and central venous (▾) pressures, and systemic vascular conductance. Right column: arterial O2 content, systemic O2 delivery, systemic a‐vO2 difference, systemic O2 extraction, and systemic O2 uptake during incremental exercise to exhaustion plotted against the relative increase in power output. Data are means ± SEM for eight subjects. * Lower than 80% of peak power, P < 0.05. Adapted from with permission of the Physiological Society and Wiley‐Blackwell.



Figure 2.

Regional distribution of cardiac output during various intensities of exercise expressed as % maximal oxygen consumption. Bar graphs on the left illustrate total cardiac output (sum of all tissue flows) and the regional distribution of cardiac output to all tissues with increasing exercise intensity. Bar graphs on the right show an expansion of regional blood flow (BF) data to all tissues except skeletal muscle tissue, to enhance appreciation of the changes in BF to the other tissue with exercise (Note the change in the BF axis for the graphs on the right). Data are estimated for human values using Rowell's text book (also estimates for humans) and our data for regional BF in pigs .



Figure 3.

Schematic drawing of an arteriole (top) and of endothelium, VSM and parenchymal cell (at the bottom) illustrating mechanisms for control of vasomotor tone and diameter. Adapted from Figure of with permission of the American Physiological Society. Neurohumoral, endothelial, and metabolic influences are detailed in the bottom part of the figure. Abbreviations: KCa, calcium‐activated K+ channel; KATP, ATP‐sensitive K+ channel; Kv, voltage‐gated K+ channel; KIR, inward rectifying K+ channel; Trp, transient receptor potential channels; O2, oxygen; ATP, adenosine triphosphate; NO, nitric oxide; TXA2, thromboxane A2 and receptor; 5HT, 5‐hydroxytryptamine and receptor; P2, purinergic type 2 receptor; M, muscarinic receptor; H1 and H2, histamine type 1 and 2 receptors; B2, bradykinin type 2 receptor; ECE, endothelin‐converting enzyme; bET‐1, big endothelin‐1; ET‐1, endothelin‐1; eNOS, endothelial nitric oxide synthase; L‐arg, L‐arginine; COX‐1, cyclooxygenase 1; CYP450, cytochrome P450; ACE, angiotensin‐converting enzyme; AI, angiotensin I; AII; angiotensin II; AT1, angiotensin type 1 receptor; AT2, angiotensin type 2 receptor; ETA, endothelin type A receptor; ETB, endothelin type B receptor; PG, prostaglandins; AA, arachidonic acid; EDHF, endothelium‐derived hyperpolarizing factor; O2, superoxide anion; VGCC, voltage‐gated calcium channels; IP, prostacyclin receptor; EETs, epoxyeicosatrienoic acids; HETEs, hydroxyeicosatetraenoic acids; H2O2, hydrogen peroxide; α1, α1‐adrenergic receptor; α2, α2‐adrenergic receptor; β2, β2‐adrenergic receptor; ACh, acetylcholine; NE, norepinephrine; NPY, neuropeptide Y; P1, purinergic type 1 receptor; and Y1, neuropeptide Y receptor.



Figure 4.

Segmental distribution of control mechanisms throughout the arterial microcirculation, redrawn from with permission of the American Physiological Society.



Figure 5.

Relationships between cardiac output (solid symbols) and two‐legged blood flow (open symbols), systemic vascular conductance (solid symbols) and two‐legged vascular conductance (open symbols), and systemic O2 delivery (solid symbols) and two‐legged O2 delivery (open symbols) versus the increases in systemic oxygen consumption during incremental exercise to exhaustion. Data are means ± SEM for six subjects. Adapted from with permission of the Physiological Society and Wiley‐Blackwell.



Figure 6.

Vasodilation of arterioles and increases in blood flow (BF) can be detected within 1 s after contraction and BF increases with a similar time course in humans. (A) A record of arteriolar diameter and tissue pO2 during single fiber stimulation showing rapid dilation and decrease in pO2. Adapted from with permission of the American Physiological Society. (B) Muscle BF response to a mild‐intensity dynamic exercise bout as measured in dogs with ultrasonic flow probes. Note the immediate increase in BF at the inititation of dynamic exercise (at the arrow). Adapted from with permission of the American Physiological Society. (C) Representative diameter records in 2A branch arterioles after single contractions for durations of 400 and 600 ms of hamster retractor muscle. Adapted from with permission of the American Physiological Society. (D) Arteriolar dilation following a 250 ms tetanic contraction of 3 to 5 fibers of hamster cremaster muscle. Graphed from data from . (E) Femoral artery BF in transition from rest to passive exercise and to voluntary, one‐legged, dynamic knee‐extensor exercise at 10 W (n = , 30 W (n = , 50 W (n = , and 70 W (n = . Voluntary exercise started at 0 time. BF was significantly greater than resting BF during passive exercise and during all time points during voluntary exercise and from passive to voluntary exercise. Adapted from Rådegran and Saltin with permission of the American Physiological Society. (F) Femoral artery BF velocity (Vmean) in transition from rest to passive exercise and to voluntary, one‐legged, dynamic knee‐extensor exercise at 50 W. Knee‐extensor force (F), intramuscular pressure (IMP), arterial (BPa) and venous (BPv) blood pressure and HR in beats per minute (BPM) are presented. Onset of voluntary exercise is indicated by an arrow on the tracing for all parameters in panel F. Adapted from Rådegran and Saltin with permission of the American Physiological Society.



Figure 7.

Results of immunoblot analysis of eNOS expression throughout arteriolar trees (each branch order) of gastrocnemius and soleus muscles of rats chronically endurance trained (ET; panels A,C,D) or chronically trained with an interval sprint training program (IST; panel B). Panels A and B present results for the gastrocnemius muscle and panel C presents eNOS expression for the soleus muscle. Panel D presents results for expression of superoxide dismutase (SOD) in feed arteries and arterioles of the soleus muscle. To clarify effects of training, results are expressed relative to protein content of arteries from sedentary rats (SED) as ET/SED or IST/SED. GFA = gastrocnemius feed artery, the first branch from the feed artery is the 1A arteriole and the branches of the 1A are subsequent 2A branches etc. WG = white portion of the gastrocnemius muscle and RG = the red portion of the gastrocnemius muscle. SFA is the soleus feed artery and the branches of the arterioles in the soleus are S1A, S2A, etc. Note that ET resulted in significant increases in eNOS protein in the arterioles feeding red muscle while IST training exhibited increased eNOS in the GFA, and WG4A and RG5A arterioles. Endurance training did not alter eNOS or SOD‐1 expression in the soleus arterioles. Endurance trained results are from McAllister et al. and those for the IST rats are from Laughlin et al. . * = value different from 1.0 with p < 0.05; + = value different from 1.0 with p < 0.1.



Figure 8.

Distribution of total CO among legs, respiratory muscles (RM), and other metabolically active tissues (skin, heart, brain, kidneys, and liver) during exercise at o2 max. Adapted from with permission of the American Physiological Society.



Figure 9.

Bar graphs of calf blood flow at rest and during maximal hyperemia (post ischemic exercise) in sedentary and trained younger and older women (top panel) and men (bottom panel). Maximal hyperemia was elicited by ischemic exercise to fatigue and measuring flow after release of the occlusion. Data are mean ±SD. *P < 0.01, **P < 0.05 versus younger subjects of corresponding gender and training status. †P < 0. 01, ‡P < 0.05 versus sedentary subjects of corresponding age and gender

. § P < 0.01, §§ P < 0.05 versus men of similar age and training status. Data taken, with permission, from Martin et al. .



Figure 10.

Arm blood flow (ABF) and a‐vO2 difference during arm cranking to exhaustion in rowers and average, fit subjects. A + L, addition of leg exercise to arm cranking in the rowers. Values are means ± SE for eight average fit subjects and seven rowers. Adapted from Volianitis et al. with permission of the American Physiological Society.



Figure 11.

Upper limb (arm) and lower limb (leg) muscle blood flow (BF) (top) and vascular conductance (bottom) from rest to peak exercise. Submaximal (SM) exercise was expressed in terms of relative exercise intensity, SM1, SM2, and SM3 (approx. 40%, 60%, and 80%, of peak power respectively). Values are mean ± SEM. *; significantly different from controls. §; significantly different from swimmers. Irrespective of the group, BF max and conductance was increased significantly from rest to peak exercise. Adapted from Walther et al. with permission of Acta Physiologica.



Figure 12.

Changes in arteriolar density do not explain changes in blood flow (BF) capacity following interval sprint training (IST) and endurance exercise training (ET). Results illustrate the effects of training on oxidative capacity (cytochrome c concentrations), BF capacity, capillary density, and arteriolar density in rat skeletal muscles. All data are expressed as a % increase above respective SED values. The IST training consisted of 10 weeks of 6 training bouts/day, 5 days/week, with each rat running 60 m/min up a 15% incline for 2.5 min with 4.5 min of rest between bouts. The ET training program consisted of 10 to 12 weeks of treadmill running at 30 m/min, 60 min/day, 5 days/week. The top row of data are for the white portion of the gastrocnemius muscle (GW), the middle row for the red portion of the gastrocnemius muscle (GR), and the bottom row of data are for the soleus muscle. These data indicate that IST increased oxidative capacity and BF capacity most in the GW muscle and these changes were correlated with increases in capillary density and small increases in arteriolar density. In contrast, ET increased oxidative capacity and BF capacity most in the soleus and GR muscle but these changes were not correlated with changes in capillary or arteriolar density in the soleus muscle. In the GR muscle, ET increases in BF capacity were similar to increases in arteriolar density. Of interest, IST increased oxidative capacity and BF capacity in the GR muscle but neither capillary nor arteriolar density were altered in this muscle tissue by IST. These data were taken from .



Figure 13.

Leg blood flow responses to graded upright cycle ergometer in younger and older healthy adults. Most results indicate that the hyperemic response to large muscle dynamic exercise is blunted in older humans. Adapted from Proctor and Parker with permission of the Microcirculatory Society.



Figure 14.

Myocardial oxygen balance in awake dogs at rest and during four incremental levels of treadmill exercise. The increase in myocardial oxygen consumption was principally accounted for by an increase in coronary blood flow with only modest contributions of increase in hematocrit and oxygen extraction. MVO2, myocardial oxygen consumption, Hct, hematocrit, ArtSO2, arterial oxygen saturation, CVSO2, coronary venous oxygen saturation. Data are from von Restorff et al. , and have been presented as mean ± SE. *P < 0.05 versus rest. Modified from with permission of the American Physiological Society.



Figure 15.

Relations between HR and left ventricular myocardial blood flow (LVMBF) at rest and during treadmill exercise in dogs 1258, 1259, 1391, 1392, , swine and horses . Data from humans were obtained principally from young healthy male subjects performing upright bicycle exercise 1492, 1493). Data from rats have been added (solid circles) to illustrate that the high LVMBF values in this species are the result of the high heart rates, so that the rat data fall close to the regression line for the human data. Modified from with permission of the American Physiological Society.



Figure 16.

Relation between M o2 and coronary venous oxygen tension (left panel) and oxygen saturation (right panel) at rest and during treadmill exercise in humans , swine , horses , and dogs . Note that exercise does not alter coronary venous oxygen tension in swine, whereas it is already reduced at low levels of exercise in dogs. Humans and horses demonstrate an intermediate oxygen tension response. During exercise oxygen saturation decreases in horses and humans (similar to dogs) whereas in swine saturation is not affected, suggesting a rightward shift of the hemoglobin‐oxygen dissociation curve during exercise in humans and horses. Also note that despite similar resting coronary venous oxygen tensions, horses and humans have higher hemoglobin‐oxygen saturations as compared to swine and dogs, consistent with the higher P50 values reported in the latter species . Data are mean ± SE. *P < 0.05 versus corresponding rest. See text for further explanation. Modified from with permission of the American Physiological Society.



Figure 17.

Coronary blood flow and hemodynamic responses to treadmill exercise in dogs. Modified from Khouri et al.. L. Circ. = left circumflex coronary artery; Cor. = coronary; Syst. = systolic; Diast. = diastolic. See text for further explanation. Modified from with permission of the American Physiological Society.



Figure 18.

Coronary pressure‐flow relation in the dog heart under conditions of maximal coronary vasodilation with intracoronary adenosine (50 μg/kg/min). Shown are the relations at rest and during three incremental levels of treadmill exercise. Note the rightward shift of the pressure‐flow relation with an increase in the zero flow pressure (Pzf) intercept. See text for further explanation. Data are from Duncker et al. and have been presented as mean ± SE. *P < 0.05 versus corresponding rest. Modified from with permission of the American Physiological Society.



Figure 19.

Graph showing a schematic drawing of the intramyocardial microvasculature (upper panel) and the extravascular forces acting on the coronary microvasculature during diastole (left lower panel) and systole (lower right panel). Abbreviations: PIM = intramyocardial pressure; PLUMEN = pressure in left ventricular lumen; PPERI = pressure in pericardial space; PPERI = pressure in pericardial space; Left ventricular lumen pressure. See text for further explanation. Modified from with permission of the American Physiological Society.



Figure 20.

Distribution of left ventricular myocardial blood flow in the dog at rest (open symbols) and during exercise (solid symbols) in the presence of intact vasomotor tone (circles) and during maximum vasodilation with intracoronary adenosine (squares). Data are from Duncker et al. and have been presented as mean ± SE. *P < 0.05 versus corresponding rest. Dot inside symbol denotes significant (P < 0.05) increase in flow produced by adenosine. Modified from with permission of the American Physiological Society.



Figure 21.

Relation between heart rate (HR) and left ventricular subendocardial to subepicardial blood flow (BF) ratio (LV ENDO/EPI) at rest and during treadmill exercise in dogs , swine 1004, 1005, 1205, 1206) and horses , measured with microspheres 15 μm in diameter. No data on transmural distribution of left ventricular BF are available in humans. Note that exercise results in a modest decrease in ENDO/EPI ratio, but this typically does not decrease below 1.0 even during heavy exercise. Also note that ENDO/EPI ratios are lower and decrease more during exercise in horses (LV E/E = −1.37•HR + 1.30) than in either swine (LV E/E = −1.17•HR + 1.38) or dogs (LV E/E = −1.06•HR + 1.41). Modified from with permission of the American Physiological Society.



Figure 22.

Relation beween myocardial oxygen consumption (MVO2) and coronary venous oxygen tension (CVPO2) in the right ventricle (RV) and the left ventricle (LV) in dogs during treadmill exercise. Note the lower levels of MVO2 and higher levels of CVPO2 in the RV compared to the LV. Data are from Hart et al. for RV data and from Gorman et al. for LV data. Data have been presented as mean ± SE. See text for further explanation. Modified from with permission of the American Physiological Society.



Figure 23.

Left panels show the effect of mixed α1‐/α2‐adrenergic receptor blockade with phentolamine on the relation between myocardial oxygen consumption (MVO2) and coronary venous oxygen tension (CVPO2) in the left ventricles of dogs and swine during treadmill exercise. Right panels show the effect of combined α1‐/α2 and β1‐/β2‐adrenergic receptor blockade with phentolamine and propranolol in swine and dogs . Data are mean ± SE. *P < 0.05 versus corresponding control or versuscorresponding phentolamine alone. See text for further explanation. Modified from with permission of the American Physiological Society.



Figure 24.

Integrative metabolic control of coronary vasomotor tone in dogs (left panels) and swine (right panels) at rest and during treadmill exercise. Shown are the effects of adenosine receptor blockade with 8‐phenyltheophylline [8PT 5 mg/kg iv ; upper panels], the effects of KATP channel blockade with glibenclamide [Glib; 50 μg/kg/min ic , or 3 mg/kg iv ] and additional adenosine receptor blockade (middle panels), and the effects of NO synthase inhibition with N‐nitro‐L‐arginine [LNA, 1.5 mg/kg ic or 20 mg/kg iv ] and additional adenosine receptor blockade and KATP channel blockade (lower panels) on the relation between myocardial oxygen consumption (M o2) and coronary venous oxygen tension (CVPO2) in the left ventricles. Data in dogs are from references . Swine data are from Merkus et al. . Data are mean ± SEM. *P < 0.05 effect of 8PT; P < 0.05 effect of glibenclamide; P < 0.05 effect of LNA. See text for further explanation. Modified from with permission of the American Physiological Society.



Figure 25.

Effects of exercise training in swine on DNA labeling in capillaries (top left panel), sprouting of new capillaries (top middle panel), % labeling of sprouts (top right panel), capillary diameter (bottom left panel), capillary density (bottom middle panel), and coronary transport reserve (CTR; bottom right panel). Data are from White et al. and have been presented as mean ± SE. *P < 0.05 versus 0 week time point (sedentary swine). See text for further explanation. Modified from with permission of the American Physiological Society.



Figure 26.

Graph summarizing the structural and functional coronary microcirculatory adaptations to chronic exercise training. ACh = acetylcholine; M = muscarinic receptor; NE = norepinephrine; α1 = α1‐adrenergic receptor. β2 = β2‐adrenergic receptor. Modified from with permission of the American Physiological Society.



Figure 27.

Indices of cerebral blood flow (BF) and oxygenation during progressive exercise expressed as % of baseline value. Arterial carbon dioxide tension (Paco2). Frontal lobe oxygenation (Sco2) determined with near‐infrared spectroscopy. The calculated average cerebral mitochondrial oxygen tension (PmitoO2). The cerebral oxygen consumption rate (CMRO2). Middle cerebral artery mean flow velocity (MCA Vmean). Cerebral BF expressed as grey matter flow (F1) and as the average flow (ISI) during cycling exercise. ISI and F1 were not determined during maximal exercise. Data from .

Courtesy of T. Seifert, the Copenhagen Muscle Center, University of Copenhagen.


Figure 28.

Linear relationships between middle cerebral artery mean flow velocity (MCA Vmean) and cardiac output ( ) at rest and during exercise. • Rest ○ Exercise. Data are means (SEM). Modified from with permission of the Physiological Society and Wiley‐Blackwell.



Figure 29.

Effects of leg muscle tensing on arterial pressure oscillations and low‐frequency power spectrum. Left: original recording. Middle: beat‐to‐beat values. Right: fast Fourier power spectrum of mean arterial pressure. The amplitude of the oscillations in arterial pressure during standing (upper panel) is attenuated during leg tensing (lower panel) with a reduction in the low‐frequency (0.07 to 0.15 Hz) power spectrum. Modified from, with permission, the Ph.D. Thesis of Pott, 2003 (1096, 1097).



Figure 30.

Regional cerebral blood flow (rCBF) by 133Xe SPECT at rest and during static handgrip exercise before and after regional blockade in 1 subject. Scale (right) represents absolute flow values. There is an increase in flow to the motor sensory area in control contraction condition (upper right) compared with rest or with contractions after axillary blockade. Premotor rCBF increased from 55 to 60 ml/100 g/liter/min (n = and motor sensor rCBF from 57 to 63 ml/liter/100 g/liter/min to both the ipsilateral and contralateral sides during handgrip exercise before, but not after, axillary blockade. Modified from Friedman et al. with permission of the American Physiological Society.



Figure 31.

Schematic description of the thermoregulatory control of skin blood flow (BF) as modified by moderately intense exercise. The relation of skin BF to internal temperature is affected, relative to resting conditions, in at least three ways by exercise: a vasoconstrictor response at the onset of dynamic exercise (A), an increase in the internal temperature threshold at which skin BF begins to increase (B), and a leveling off, or plateau, in skin BF above an internal temperature of 38°C at a level well below maximal (C). Exercise exerts these effects through the vasoconstrictor system for the initial vasoconstriction and through inhibiting the active vasodilator system for the increased threshold and for the plateau. At rest, the plateau only occurs as skin BF approaches maximal vasodilation. Modified from Gonzalez‐Alonso et al. 2008 with permission of the Physiological Society and Wiley‐Blackwell.



Figure 32.

Heterogeneity of blood flow (BF) to various bone tissue in anesthetized dogs. Values below the bars reflect the number of observations for the mean value illustrated. Epiphysis is from the femur inferior epiphysis; “diaphysis represents 28 samples of femur diaphysis and 8 samples of humerus diaphysis.” BFs from the two diaphyseal regions were not significantly different (P > 0.05; “red marrow” is from the femur medulla, and “yellow marrow” represents samples from the medullas of the tibia and mandible. Analysis of variance and Tukey's test indicate the following significant differences: sternum, rib, ilium, and epiphysis > diaphysis and yellow marrow; sternum > rib; red and yellow marrow > diaphysis; red marrow > yellow marrow. Modified from Gross et al. 1979 with permission of the American Physiological Society.



Figure 33.

Relationship between splanchnic blood flow (BF) and exercise intensity (expressed as percent o2 max) at normal temperature (25.6°C, solid line) and during heat stress (43.3°C, dashed line). The reduction of splanchnic BF during exercise is proportional to the relative exercise intensity (% o2 max) and is further reduced by environmental heat stress resulting in ∼20% greater reductions in splanchnic BF at a given relative exercise intensity. Modified from Rowell, 1993 which was adapted from Rowell 1965 with permission of the American Physiological Society.



Figure 34.

Exercise‐induced reductions in both splanchnic blood flow (BF) and splanchnic blood volume are nonuniformly distributed across the splanchnic organs. (A) Nonuniform changes in splanchnic BF during and 2 minutes after termination of cycling. The decreases in total splanchnic BF (SBF) (top panel) during moderate cycling exercise involve greater reductions in celiac artery BF (dotted line; middle panel) than in the superior mesenteric artery BF (solid line; middle panel) owing to differential increases in artery resistance (R) in the celiac (dotted line) and superior mesenteric artery (solid line) vascular beds (bottom panel). For both BF and R data in the middle and bottom panels, respectively; continuous line = superior mesenteric artery and the dashed line = the celiac artery. * = different from the preceding value, P < 0.05; # = different from rest, P < 0.05. Data from Perko et al. with permission of the American Physiological Society. (B) Nonuniform changes in splanchnic blood volume. Splanchnic blood volume redistribution during zero‐load (ZLC) and graded cycling exercise (50%‐100% maximal exercise) and 5 min of recovery from exercise. Note that the volume redistribution involves a greater reduction of splenic than hepatic and renal blood volume and that bowel blood volume did not significantly change from baseline values during or following exercise. Values are % basal volume, mean ± SE. Data taken from Table of Flamm et al. 1990 .



Figure 35.

Exercise training alters the redistribution of blood flow (BF) to kidney and splanchnic tissues during exercise. Top panel: baseline/resting BF values represent sedentary rats standing quietly on the treadmill. Middle panel: exercise BF values for sedentary rats at 15 min of treadmill running 30 m/min. Bottom panel: exercise BF values for exercise trained rats at 15 min of treadmill running 30 m/min. Baseline BFs were similar between sedentary and trained rats. As shown in the middle panel, exercise caused BF reductions to all organs, except the liver in sedentary animals. In contrast, results presented in the bottom panel indicate that trained animals exhibit an attenuated reduction of splanchnic and renal BFs during acute exercise. Values are mean ± SE. Liver BF represents hepatic artery BF only. Data are from Armstrong and Laughlin and McAllister .



Figure 36.

Graded cycling exercise markedly increases renal vascular resistance thereby reducing renal blood flow (inferred from reduced renal venous outflow). Workloads were 30%, 60%, and 80% to 90% o2 max. Values are mean ± SE. The *, **, and *** indicate values significantly different from rest, with p < 0.05, 0.01, and 0.001, respectively. Adapted from Tidgren et al. with permission of the American Physiological Society

.



Figure 37.

Changes in pulmonary artery pressure (PAP, open circles in left panels), left atrial pressure (solid circles in left panels) and pulmonary vascular resistance (PVR, right panels) as a function of estimated exercise intensity in dogs , swine , sheep , humans , and horses . Note that PVR is inversely related to body weight (dogs ∼15 kg; swine and sheep ∼25 kg; humans ∼75 kg; and horses ∼400 kg). Exercise intensity is calculated based on heart rate (dogs, sheep, and horses) and/or body oxygen consumption (swine and humans). From Merkus et al. with permission of Elsevier.



Figure 38.

Integrated endothelial control of pulmonary vascular tone in swine (554). The vasodilator effect of ETA/ETB receptor blockade with tezosentan was enhanced by prior inhibition of eNOS (middle panels), but not by COX inhibition (bottom panels) at rest and during exercise in swine. These findings indicate that NO limits the influence of ET on the pulmonary vasculature. B o2: body oxygen consumption; PAP: pulmonary artery pressure; PVR: pulmonary vascular resistance. *P < 0.05 versus control, †P < 0.05 effect of tezosentan different after eNOS/COX inhibition. Adapted from Merkus et al. with permission of Elsevier.



Figure 39.

Autonomic control of pulmonary vascular tone in swine . α‐Adrenoceptor blockade results in vasodilation (top), whereas β‐adrenoceptor blockade results in vasoconstriction at rest and during exercise (middle). Hence, the sympathetic nervous system exerts no net effect on pulmonary vascular tone. M‐receptor blockade results in pulmonary vasodilation at rest that wanes with increasing exercise intensity (bottom), suggesting that withdrawal of the vasoconstrictor influence exerted by the parasympathetic nervous system contributes to exercise‐induced pulmonary vasodilation. B o2: body oxygen consumption; PAP: pulmonary artery pressure; PVR: pulmonary vascular resistance. *P < 0.05 versus control. From Merkus et al. with permission of Elsevier.



Figure 40.

Schematic regulation of the pulmonary vascular tone at rest and during exercise in the healthy pulmonary circulation. NO: nitric oxide; PGI2: prostacyclin; βAR: β adrenoceptor; MR: muscarinic receptor; Ado: adenosine; ANP: atrial natriuretic peptide; KATP, KCa2+ and KV: ATP, Ca2+ and voltage‐sensitive K‐channels; ETA: endothelin A receptor; ETB: endothelin B receptor; αAR: α‐adrenoceptor; Ang‐II: angiotensin II; TXA2: thromboxane A2; 5‐HT: 5‐hydroxytryptamine; PDE5: phosphodiesterase 5; ROS: reactive oxygen species; +: significant influence; −: no influence; ?: probable influence (not yet proven). See text for further explanations. From Merkus et al. with permission of Elsevier.

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M. Harold Laughlin, Michael J. Davis, Niels H. Secher, Johannes J. van Lieshout, Arturo A. Arce‐Esquivel, Grant H. Simmons, Shawn B. Bender, Jaume Padilla, Robert J. Bache, Daphne Merkus, Dirk J. Duncker. Peripheral Circulation. Compr Physiol 2012, 2: 321-447. doi: 10.1002/cphy.c100048