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RESPIRATORY MECHANICS AND GAS EXCHANGE  103
                          average slope, or compliance, that characterize the
                          loop and have significant effects on their counterparts
                          in Fig. 4.7.
                            Insufficient surfactant levels can occur in premature
                          neonates whose alveolar cells are not mature enough to
                          produce sufficient quantities, a condition leading to respi-
                          ratory distress syndrome, also called hyaline membrane
                               2
                          disease. Instilling liquid mixtures, containing either nat-
                          ural or man-made surfactants, directly into airways via
                          the trachea has developed from early work in animal
                               9
                          models into an important clinical tool called surfactant
                          replacement therapy.  The movement of these liquid
                          boluses through the network relies on several mecha-  FIGURE 4.10  Surface area  A versus surface
                                                                                        s
                          nisms, including air-blown liquid plug flow dynamics,  tension  s for a cycled interface containing lung
                          gravity, and surface tension and its gradients. 4,10,18  surfactant. There is a hysteresis area and mean slope
                                                                   or compliance to the curve; compare to Fig. 4.7.
              4.5 VENTILATION, PERFUSION, AND LIMITS

                          The lung differs from many other organs in its combina-
                          tion of gas-phase and liquid-phase constituency. Because
                          the densities of these two phases are so different, grav-
                          ity plays an important role in determining regional lung
                          behavior, both for gas ventilation and for blood perfu-
                          sion. In an upright adult, the lower lung is compressed
                          by the weight of the upper lung and this puts the lower
                          lung’s alveoli on a more compliant portion of their
                          regional P-V curve; see Fig. 4.7. Thus inhaled gas tends
                          to be directed preferentially to the lower lung regions,

                          and the regional  alveolar ventilation V A  decreases in a
                          graded fashion moving upward in the gravity field.

                            Blood flow Q  is also preferentially directed toward
                          the lower lung, but for different reasons. The blood pres-
                          sure in pulmonary arteries,  P , and veins,  P , sees a
                                                           v
                                                a
                          hydrostatic pressure gradient in the upright lung. These
                          vessels are imbedded within the lung’s structure, so they
                          are surrounded by alveolar gas pressure, P , which is
                                                         A
                          essentially uniform in the gravity field since its density
                          is negligible. Thus there is a region called Zone III in the
                          upper lung where P < P . This difference in pressures
                                            A
                                        a
                          squeezes the capillaries essentially shut and there is rel-
                          atively little pulmonary blood flow there. In the lower
                          lung called Zone I, the hydrostatic effect is large enough
                          to keep P > P > P and blood flow is proportional to
                                    v
                                 a
                                        A
                          the arterial-venous pressure difference,  P − P . In
                                                         a
                                                              v
                          between these two zones is Zone II where P > P > P .
                                                         a
                                                            A
                                                                v
                          Here there will be some length of the vessel that is nei-
                          ther fully closed nor fully opened. It is partially col-
                          lapsed into more of a flattened, oval shape. The physics
                          of the flow, called the  vascular waterfall 29  or  choked
                                           23
                          flow 30  or flow limitation , dictates that Q is no longer  FIGURE 4.11 Choked flow through a flexible
                                                                    tube. (a) upstream pressure P , downstream pressure
                                                                                    u
                          dependent on the downstream pressure, but is primarily  P , external pressure P , flow F, pressure P, cross-
                                                                     d
                                                                                 ext
                          determined by the pressure difference P − P .  sectional area A. (b) A/A versus transmural pressure
                                                      a
                                                         A
                                                                                  0
                            Figure 4.11a shows this interesting type of flexible  with shapes indicated. (c) Flow versus pressure drop,
                          tube configuration and flow limitation phenomena where  assuming P is decreased and P is fixed.
                                                                                      u
                                                                          d
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