physiology of decompression physiology of decompression the physiology of decompression is the aspect of physiology which is affected by exposure to large changes in ambient pressure, and involves a complex interaction of gas solubility, partial pressures and concentration gradients, diffusion, bulk transport and bubble mechanics in living tissues. gas is breathed at ambient pressure, and some of this gas dissolves into the blood and other fluids. inert gas continues to be taken up until the gas dissolved in the tissues is in a state of equilibrium with the gas in the lungs, (see: "saturation diving"), or the ambient pressure is reduced until the inert gases dissolved in the tissues are at a higher concentration than the equilibrium state, and start diffusing out again. scuba diver decompressing at a planned stop during ascent from a dive not to be confused with human physiology of underwater diving or decompression sickness. the absorption of gases in liquids depends on the solubility of the specific gas in the specific liquid, the concentration of gas, customarily expressed as partial pressure, and temperature. in the study of decompression theory the behaviour of gases dissolved in the body tissues is investigated and modeled for variations of pressure over time. once dissolved, distribution of the dissolved gas is by perfusion, where the solvent (blood) is circulated around the diver's body, and by diffusion, where dissolved gas can spread to local regions of lower concentration when there is no bulk flow of the solvent. given sufficient time at a specific partial pressure in the breathing gas, the concentration in the tissues will stabilise, or saturate, at a rate depending on the local solubility, diffusion rate and perfusion. if the concentration of the inert gas in the breathing gas is reduced below that of any of the tissues, there will be a tendency for gas to return from the tissues to the breathing gas. this is known as outgassing, and occurs during decompression, when the reduction in ambient pressure or a change of breathing gas reduces the partial pressure of the inert gas in the lungs. the combined concentrations of gases in any given tissue will depend on the history of pressure and gas composition. under equilibrium conditions, the total concentration of dissolved gases will be less than the ambient pressure, as oxygen is metabolised in the tissues, and the carbon dioxide produced is much more soluble. however, during a reduction in ambient pressure, the rate of pressure reduction may exceed the rate at which gas can be eliminated by diffusion and perfusion, and if the concentration gets too high, it may reach a stage where bubble formation can occur in the supersaturated tissues. when the pressure of gases in a bubble exceed the combined external pressures of ambient pressure and the surface tension from the bubble - liquid interface, the bubbles will grow, and this growth can cause damage to tissues. symptoms caused by this damage are known as decompression sickness. the actual rates of diffusion and perfusion, and the solubility of gases in specific tissues are not generally known, and vary considerably. however mathematical models have been proposed which approximate the real situation to a greater or lesser extent, and these models are used to predict whether symptomatic bubble formation is likely to occur for a given pressure exposure profile. efficient decompression requires the diver to ascend fast enough to establish as high a decompression gradient, in as many tissues, as safely possible, without provoking the development of symptomatic bubbles. this is facilitated by the highest acceptably safe oxygen partial pressure in the breathing gas, and avoiding gas changes that could cause counterdiffusion bubble formation or growth. the development of schedules that are both safe and efficient has been complicated by the large number of variables and uncertainties, including personal variation in response under varying environmental conditions and workload. solubility solubility is the property of a gas, liquid or solid substance (the solute) to be held homogeneously dispersed as molecules or ions in a liquid or solid medium (the solvent). in decompression theory the solubility of gases in liquids is of primary importance, as it is the formation of bubbles from these gases that causes decompression sickness. solubility of gases in liquids is influenced by three main factors: the nature of the solvent liquid and the solute temperature (gases are less soluble in water but may be more soluble in organic solvents, at higher temperatures.) pressure (solubility of a gas in a liquid is proportional to the partial pressure of the gas on the liquid – henry's law, the presence of other solutes in the solvent can also influence solubility. body tissues include aqueous and lipid components in varying ratios, and the solubility of the gases involved in decompression in these tissues will vary depending on their composition. solubility of gases at 37 °c gas molecular weight lipid/water solubility ratio hydrogen 2 3.1 helium 4 1.7 neon 20 2.07 nitrogen 28 5.2 diffusion diffusion is the movement of molecules or ions in a medium when there is no gross mass flow of the medium, and can occur in gases, liquids or solids, or any combination. diffusion is driven by the kinetic energy of the diffusing molecules – it is faster in gases and slower in solids when compared with liquids due to the variation in distance between collisions, and diffusion is faster when the temperature is higher as the average energy of the molecules is greater. diffusion is also faster in smaller, lighter molecules of which helium is the extreme example. diffusivity of helium is 2.65 times faster than nitrogen. the partial pressure gradient, also known as the concentration gradient, can be used as a model for the driving mechanism of diffusion. the partial pressure gradient is the variation of partial pressure (or more accurately, the concentration) of the solute (dissolved gas) from one point to another in the solvent. the solute molecules will randomly collide with the other molecules present, and tend over time to spread out until the distribution is statistically uniform. this has the effect that molecules will diffuse from regions of higher concentration (partial pressure) to regions of lower concentration, and the rate of diffusion is proportional to the rate of change of the concentration. tissues in which an inert gas is more soluble will eventually develop a higher dissolved gas content than tissues where the gas is less soluble. inert gas uptake (ingassing) in this context, inert gas refers to a gas which is not metabolically active. atmospheric nitrogen (n2) is the most common example, and helium (he) is the other inert gas commonly used in breathing mixtures for divers. atmospheric nitrogen has a partial pressure of approximately 0.78 bar at sea level. air in the alveoli of the lungs is diluted by saturated water vapour (h2o) and carbon dioxide (co2), a metabolic product given off by the blood, and contains less oxygen (o2) than atmospheric air as some of it is taken up by the blood for metabolic use. the resulting partial pressure of nitrogen is about 0,758 bar. at atmospheric pressure the body tissues are therefore normally saturated with nitrogen at 0.758 bar (569 mmhg). at increased ambient pressures due to depth or habitat pressurisation, a diver's lungs are filled with breathing gas at the increased pressure, and the partial pressures of the constituent gases will be increased proportionately. for example: at 10 meters sea water (msw) the partial pressure of nitrogen in air will be 1.58 bar. the inert gases from the breathing gas in the lungs diffuse into blood in the alveolar capillaries ("move down the pressure gradient") and are distributed around the body by the systemic circulation in the process known as perfusion. perfusion perfusion is the mass flow of blood through the tissues. dissolved materials are transported in the blood much faster than they would be distributed by diffusion alone (order of minutes compared to hours). the dissolved gas in the alveolar blood is transported to the body tissues by the blood circulation. there it diffuses through the cell membranes and into the tissues, where it may eventually reach equilibrium. the greater the blood supply to a tissue, the faster it will reach equilibrium with gas at the new partial pressure. saturation and supersaturation if the supply of gas to a solvent is unlimited, the gas will diffuse into the solvent until there is so much dissolved that equilibrium is reached and the amount diffusing back out is equal to the amount diffusing in. this is called saturation. if the external partial pressure of the gas (in the lungs) is then reduced, more gas will diffuse out than in. this is a condition known as supersaturation. the gas will not necessarily form bubbles in the solvent at this stage, but supersaturation is necessary for bubble growth. a supersaturated solution of gases in a tissue may form bubbles if suitable nucleation sites exist. supersaturation may be defined as a sum of all gas partial pressures in the liquid which exceeds the ambient pressure in the liquid. tissue half times if an exponential uptake of gas is assumed, which is a good approximation of experimental values for diffusion in non-living homogenous materials, half time of a tissue is the time it takes for the tissue to take up or release 50% of the difference in dissolved gas capacity at a changed partial pressure. for each consecutive half time the tissue will take up or release half again of the cumulative difference in the sequence ½, ¾, 7/8, 15/16, 31/32, 63/64 etc. the number of half times chosen to assume full saturation depends on the decompression model, and typically ranges from 4 (93.75%) to 6 (98.44%). tissue compartment half times used in decompression modelling range from 1 minute to at least 720 minutes. for example: a 5 minute tissue will be 50% saturated in 5 minutes, 75% in 10 minutes, 87.5% in 15 minutes and for practical purposes, saturated in about 30 minutes (98.44% saturated at 6 half times) a specific tissue compartment will have different half times for gases with different solubilities and diffusion rates. this model may not adequately describe the dynamics of outgassing if gas phase bubbles are present. outgassing of tissues gas remains dissolved in the tissues until the partial pressure of that gas in the lungs is reduced sufficiently to cause a concentration gradient with the blood at a lower concentration than the relevant tissues. a lowered partial pressure in the lungs will result in more gas diffusing out of the blood into the lung gas and less from the lung gas into the blood. a similar situation occurs between the blood and each tissue. as the concentration in the blood drops below the concentration in the adjacent tissue, the gas will diffuse out of the tissue into the blood, and will then be transported back to the lungs where it will diffuse into the lung gas and then be eliminated by exhalation. if the ambient pressure reduction is limited, this desaturation will take place in the dissolved phase, but if the ambient pressure is lowered sufficiently, bubbles may form and grow, both in blood and other supersaturated tissues. when the gas in a tissue is at a concentration where more diffuses out than in, the tissue is said to be supersaturated with that gas relative to the surrounding tissues. supersaturation can also be defined as when the combined partial pressures of gases dissolved in a tissue exceeds the total ambient pressure on the tissue, and there is a theoretical possibility of bubble formation or growth. inherent unsaturation there is a metabolic reduction of total gas pressure in the tissues. the sum of partial pressures of the gas that the diver breathes must necessarily balance with the sum of partial pressures in the lung gas. in the alveoli the gas has been humidified by a partial pressure of approximately 63 mbar (47 mmhg) and has gained about 55 mbar (41 mmhg) carbon dioxide from the venous blood. oxygen has also diffused into the arterial blood, reducing the partial pressure of oxygen in the alveoli by about 67 mbar(50 mmhg) as the total pressure in the alveoli must balance with the ambient pressure, this dilution results in an effective partial pressure of nitrogen of