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INTRODUCTIONThe primary function of the respiratory system is to oxygenate blood and eliminate carbon dioxide, which requires that blood come into virtual contact with fresh air to facilitate diffusion of respiratory gases between blood and gas. This process occurs in the lung alveoli, where blood flowing through alveolar wall capillaries is separated from alveolar gas by an extremely thin membrane of flattened endothelial and epithelial cells, across which respiratory gases diffuse and equilibrate. Blood flow through the lung is unidirectional via a continuous vascular path, along which venous blood absorbs oxygen from and loses CO2 to inspired gas. The path for airflow, in contrast, reaches a dead end at the alveolar walls; as such, the alveolar space must be ventilated tidally, with inflow of fresh gas and outflow of alveolar gas alternating periodically at the respiratory rate (RR). To achieve an enormous alveolar surface area (typically 70 m2) for blood-gas diffusion within the modest volume of a thoracic cavity (typically 7 L), nature has distributed both blood flow and ventilation among millions of tiny alveoli through multigenerational branching of both pulmonary arteries and bronchial airways. As a consequence of variations in tube lengths and calibers along these pathways, and of the effects of gravity, tidal pressure fluctuations, and anatomic constraints from the chest wall, there is variation among alveoli in their relative ventilations and perfusions. Not surprisingly, for the lung to be most efficient in exchanging gas, the fresh gas ventilation of a given alveolus must be matched to its perfusion.For the respiratory system to succeed in oxygenating blood and eliminating carbon dioxide, it must be able to ventilate the lung tidally to freshen alveolar gas; it must provide for perfusion of the individual alveolus in a manner proportional to its ventilation; and it must allow for adequate diffusion of respiratory gases between alveolar gas and capillary blood. Furthermore, it must be able to accommodate severalfold increases in the demand for oxygen uptake or CO2 elimination imposed by metabolic needs or acid-base derangement. Given these multiple requirements for normal operation, it is not surprising that many diseases disturb respiratory function. Here, we consider in greater detail the physiologic determinants of lung ventilation and perfusion, and how their matching distributions and rapid gas diffusion allow for normal gas exchange. We also discuss how common diseases derange these normal functions, and thereby impair gas exchange—or at least raise the work of the respiratory muscles or heart to maintain adequate respiratory function.VENTILATIONIt is useful to think about the respiratory system as having three independently functioning components—the lung including its airways, the neuromuscular system, and the chest wall; the latter includes everything that is not lung or active neuromuscular system. As such, the mass of the respiratory muscles is part of the chest wall, while the force they generate is part of the neuromuscular system; the abdomen (especially an obese abdomen) and the heart (especially an enlarged heart) are, for these purposes, part of the chest wall. Each of these three components has mechanical properties that relate to its enclosed volume, or in the case of the neuromuscular system, the respiratory system volume at which it is operating, and to the rate of change of its volume (i.e., flow).Volume-Related Mechanical Properties-StaticsFigure A-1 shows the volume-related properties of each component of the respiratory system. Due both to surface tension at the air-liquid interface between alveolar wall lining fluid and alveolar gas and to elastic recoil of the lung tissue itself, the lung requires a positive transmural pressure difference between alveolar gas and its pleural surface to stay inflated; this difference is called the elastic recoil pressure of the lung, and it increases with lung volume. Importantly, the lung becomes rather stiff at high lung volumes, so that relatively small volume changes are accompanied by large changes in transpulmonary pressure; in contrast, the lung is compliant at lower lung volumes, including those at which tidal breathing normally occurs. Note that at zero inflation pressure, even normal lungs retain some air in the alveoli. This occurs because the small peripheral airways of the lung are tethered open by radially outward pull from inflated lung parenchyma attached to adventitia; as the lung deflates during exhalation, those small airways are pulled open progressively less, and eventually they close, trapping some gas in the alveoli. This effect can be exaggerated with age and especially with obstructive airways diseases, resulting in gas trapping at quite large lung volumes.
Figure A-1 . Pressure-volume curves of the isolated lung, isolated chest wall, combined respiratory system, inspiratory muscles, and expiratory muscles. FRC, functional residual capacity; RV, residual volume; TLC, total lung capacity.
The elastic behavior of the passive chest wall (i.e., in the absence of neuromuscular activation) differs markedly from that of the lung. Whereas the lung tends toward full deflation with no distending (transmural) pressure, the chest wall encloses a large volume when pleural pressure equals body surface (atmospheric) pressure. Furthermore, the chest wall is compliant at high enclosed volumes, readily expanding even further in response to increases in transmural pressure. The chest wall also remains compliant at small negative transmural pressures (i.e., when pleural pressure falls slightly below atmospheric pressure), but as the volume enclosed by the chest wall becomes quite small in response to large negative transmural pressures, the passive chest wall becomes stiff due to squeezing together of ribs and intercostal muscles, diaphragm stretch, displacement of abdominal contents, and straining of ligaments and bony articulations. Under normal circumstances, the lung and the passive chest wall enclose essentially the same volume, the only difference between these being the volumes of the pleural fluid and of the lung parenchyma (both quite small). As such, and because the lung and chest wall function in mechanical series, the pressure required to displace the passive respiratory system (lungs + chest wall) at any volume is simply the sum of the elastic recoil pressure of the lungs and the transmural pressure across the chest wall. When plotted against respiratory system volume, this relationship assumes a sigmoid shape, exhibiting stiffness at high lung volumes (imparted by the lung), stiffness at low lung volumes (imparted by the chest wall, or sometimes by airway closure), and compliance in the middle range of lung volumes. There is also a passive resting point of the respiratory system, attained when alveolar gas pressure equals body surface pressure (i.e., the transrespiratory system pressure is zero). At this volume [called functional residual capacity (FRC)], the outward recoil of the chest wall is balanced exactly by the inward recoil of the lung. As these recoils are transmitted through the pleural fluid, the latter is pulled both outward and inward simultaneously at FRC, and, thus, its pressure falls below atmospheric pressure (typically, –5 cmH2O).The normal passive respiratory system would equilibrate at FRC and remain there were it not for the actions of respiratory muscles. The inspiratory muscles act on the chest wall to generate the equivalent of positive pressure across the lungs and passive chest wall, while the expiratory muscles generate the equivalent of negative transrespiratory pressure. The maximal pressures these sets of muscles can generate varies with the lung volume at which they operate, due to length-tension relationships in striated muscle sarcomeres and to changes in mechanical advantage as the angles of insertion change with lung volume (Fig. A-1). Nonetheless, under normal conditions the respiratory muscles are substantially “overpowered” for their roles, and generate more than adequate force to drive the respiratory system to its stiffness extremes, as determined by the lung [total lung capacity (TLC)] or chest wall or airway closure [residual volume (RV)]; importantly, the latter always prevents the adult lung from emptying completely under normal circumstances. The excursion between full and minimal lung inflation is called vital capacity (VC; Fig. A-2), and is readily seen to be the difference between volumes at two unrelated stiffness extremes—one determined by the lung (TLC) and the other determined by the chest wall or airways (RV). Thus, although VC is easy to measure (see below), it tells one little about the intrinsic properties of the respiratory system. It is much more useful, as we shall see, for the clinician to know TLC and RV individually.
Fig. A-2 . Spirogram demonstrating a slow vital capacity maneuver and various lung volumes.Flow-Related Mechanical Properties—DynamicsThe passive chest wall and active neuromuscular system do exhibit mechanical behaviors related to the rate of change of volume, but these become quantitatively important only at markedly supraphysiologic breathing frequencies (e.g., during high-frequency mechanical ventilation), and, thus, we shall not address these here. In contrast, the dynamic airflow properties of the lung substantially determine the ability to ventilate and contribute importantly to the work of breathing, and are often deranged by disease. Understanding these properties is, therefore, well worthwhile.As with flow of any fluid (gas or liquid) in any tube, maintenance of airflow within the pulmonary airways requires a pressure gradient that falls along the direction of flow, the magnitude of which is determined by the flow rate and the frictional resistance to flow. During quiet tidal breathing, the pressure gradients driving inspiratory or expiratory flow are small owing to the very low frictional resistance of normal pulmonary airways (normally <2 cmH2OL per second). However, during rapid exhalation another phenomenon reduces flow below that which would have been expected were frictional resistance the only impediment to flow. This is called dynamic airflow limitation, and it occurs because bronchial airways through air exhaled are collapsible rather than rigid (Fig. A-3). An important anatomic feature of pulmonary its treelike branching structure. While individual in each successive generation, from most proximal (trachea) distal (respiratory bronchioles), smaller those parent their number increases exponentially such summed cross-sectional area becomes very large toward lung periphery. Because (volume time) constant along airway tree, velocity (flow area) much greater central peripheral airways. During exhalation, gas leaving alveoli must therefore gain as proceedsmouth. The energy required for this “convective” acceleration drawn component manifested localpressure, thereby reducing intraluminal pressure (the Bernoulli effect), transmural size A-3), If one tries exhale more forcefully, further further, resulting no net increase Under these circumstances, has reached maximum possible value, or limit. Lungs normally exhibit limitation. value related density, cross-section distensibility, elastic recoil lung, loss flow-limiting site. normal conditions, maximal expiratory falls with volume A-4), due primarily dependence on A-1). In fibrosis, increased at any volume, and, thus, relatively elevated when considered relation volume. Conversely, emphysema, reduced, a principal mechanism by flows fall. Diseases narrow lumen asthma chronic bronchitis, cause excessive collapsibility, like tracheomalacia, also reduce flow.
Fig. A-3. Luminal versus relationship. Transmural represents difference across wall inside outside.
Fig. A-4. Flow-volume loops. A. Normal. B. Airflow obstruction. C. Fixed RV, residual volume; TLC, total capacity.The effect acts inspiration, but negative pleural pressures inspiration lower outside airways, increasing promoting expansion. Thus, inspiratory limitation seldom diffuse disease. extrathoracic narrowing (as tracheal adenoma post-tracheostomy stricture) can lead A-4).The importance distensibility upstream (lung forced exhalation) easily be appreciated sniffing one’s nose low, medium, substantial effort. keeps nostrils relaxed, low medium effort raises somewhat, inhaling even harder will likely not nasal but, rather, just collapse nares, manifestation One airflow, however, flaring using alae nasi muscles. nostril (reducing given nares) stiffens walls response pressure). Springlike strips sometimes used football players same effect. patients obstructive sleep apnea (OSA), narrowed excessively compliant pharyngeal collapses generated (which why an upper respiratory infection often worsens OSA). Increasing driving phenomena intrapharyngeal positive positive, preventing Inspiratory pharynx OSA closely parallels lung.Work BreathingIn health, change–related) (flow-related) loads overcome ventilate lungs rest small, work muscles minimal. breathing considerably, either requirement substantially ventilation, abnormally mechanical load, both. As discussed below, rate ventilation set need eliminate carbon dioxide, exercise (sometimes 20-fold) metabolic acidosis compensatory response. Naturally, elasticity system both depth frequency tidal breaths, while load ventilation. A modest efficiently achieved rate, ventilatory level exercise. At high levels exercise, deep persists, increases. pattern chosen controller minimizes breathing.Work disease compliance airflow. former commonly diseases parenchyma (interstitial processes alveolar filling edema pneumonia, resection), latter asthma, cystic fibrosis. Furthermore, severe obstruction functionally leading hyperinflation. scenario, slowed may insufficient allow complete phase breathing; result, “functional capacity” next breath inhaled static FRC. With repetition incomplete exhalations breath, operating FRC dynamically elevated, approaches TLC. volumes, less increased. hyperinflation accompanies causes sense difficulty in—even though pathophysiologic abnormality root obstruction.Adequacy VentilationAs noted above, control sets responds chemical signals, including arterial dioxide oxygen tensions blood pH, volitional needs, inhale deeply before playing long phrase trumpet. Here, we focus relationship between elimination.At end conducting filled had mouth stopped. ensuing inhalation, fresh immediately enters tree mouth, first entering start inhalation left alveoli. such, does enter until inspired. dead space. Quiet volumes space introduces into all; part inspired (VT) (VD) Importantly, if some delivered receives flow, cannot contribute exchange, occur portion embolus. minute (VE =VT x RR) includes (VD =VD (VA =[VT-VD] Carbon elimination equal VA times CO2 fraction (essentially zero) (typically ~5.6%, after correcting humidification air, corresponding 40 mmHg). steady state, production divided Because, equal, strives maintain PCO2 (PaCO2) ~40 mmHg, adequacy reflected PaCO2. PaCO2 hyperventilation present, exceeds then hypoventilation present. Ventilatory failure characterized extreme hypoventilation.As consequence uptake capillary blood, tension gas. (determined body’s consumption) average ratio, “respiratory quotient” (R =Vco2/Vo2), depends largely fuel being metabolized. For typical merican diet, R usually around 0.85, absorbed excreted. Together, estimation tension, according following relationship, known equation:PaO2 =FIO2 (Pbar – PH2O) RThe equation highlights influences (FIO2), barometric (Pbar), vapor water (PH2O =47 mmHg 37°C) addition PACO2) determining PAO2. implication hypoxemia rarely pure sea air. potential induce otherwise Pbar altitude.Gas ExchangeDiffusionFor tissues, pass diffusing membrane. aggregate membrane highly optimized process, surface minimal thickness. Diffusion so efficient human circumstances hemoglobin fully saturated time red cell traveled one-third length capillary. ordinarily limited amount transiting capillaries how rapidly membrane; said “perfusion limited.” equilibrates alveolus essentially rare diffusion-limited, altitude high-performance athletes exerting interstitial thickened remain perfused.Ventilation-Perfusion HeterogeneityAs exchange efficient, should matched perfusion accompanying millions Due differential effects gravity mechanics throughout differences vascular architecture among various paths, there minor heterogeneity lung; V Q particularly marked Two examples (1) unperfused embolus, physiologic “wasted” exchange; (2) nonventilated condition “shunt.” allows venous unaltered; mixed oxygenated other well-ventilated units, shunted disproportionately lowers PaO2, nonlinear content PO2 A-5). refractory supplemental oxygen. raising FIO2 alveoli, PAO2 ventilated exiting units slightly already nearly solubility plasma quite small.
Fig. A-5. Influence vs oxygenation shunt heterogeneity. Partial (mmHg) saturations shown affected blood.More occurring two above widening distribution></div></div><footer><div
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