Gas exchange

Diffusion and surface area The exchange of gases occurs as a result of diffusion down a concentration gradient. Gas molecules move from a region in which they are at high concentration to one in which they are at low concentration. Diffusion is a passive process, meaning that no energy is required to power the transport, and it follows Fick's law: :J = -D \frac{d \varphi}{d x} In relation to a typical biological system, where two compartments ('inside' and 'outside'), are separated by a membrane barrier, and where a gas is allowed to spontaneously diffuse down its concentration gradient: • J is the flux, the amount of gas diffusing per unit area of membrane per unit time. Note that this is already scaled for the area of the membrane. • D is the diffusion coefficient, which will differ from gas to gas, and from membrane to membrane, according to the size of the gas molecule in question, and the nature of the membrane itself (particularly its viscosity, temperature and hydrophobicity). • φ is the concentration of the gas. • x is the position across the thickness of the membrane. • dφ/dx is therefore the concentration gradient across the membrane. If the two compartments are individually well-mixed, then this is simplifies to the difference in concentration of the gas between the inside and outside compartments divided by the thickness of the membrane. • The negative sign indicates that the diffusion is always in the direction that - over time - will destroy the concentration gradient, i.e. the gas moves from high concentration to low concentration until eventually the inside and outside compartments reach equilibrium. Gases must first dissolve in a liquid in order to diffuse across a membrane, so all biological gas exchange systems require a moist environment. In general, the higher the concentration gradient across the gas-exchanging surface, the faster the rate of diffusion across it. Conversely, the thinner the gas-exchanging surface (for the same concentration difference), the faster the gases will diffuse across it. In the equation above, J is the flux expressed per unit area, so increasing the area will make no difference to its value. However, an increase in the available surface area, will increase the amount of gas that can diffuse in a given time. but in larger organisms such as roundworms (Nematoda) the equivalent exchange surface - the cuticle - is substantially thicker at 0.5 μm. Interaction with circulatory systems In multicellular organisms therefore, specialised respiratory organs such as gills or lungs are often used to provide the additional surface area for the required rate of gas exchange with the external environment. However the distances between the gas exchanger and the deeper tissues are often too great for diffusion to meet gaseous requirements of these tissues. The gas exchangers are therefore frequently coupled to gas-distributing circulatory systems, which transport the gases evenly to all the body tissues regardless of their distance from the gas exchanger. Some multicellular organisms such as flatworms (Platyhelminthes) are relatively large but very thin, allowing their outer body surface to act as a gas exchange surface without the need for a specialised gas exchange organ. Flatworms therefore lack gills or lungs, and also lack a circulatory system. Other multicellular organisms such as sponges (Porifera) have an inherently high surface area, because they are very porous and/or branched. Sponges do not require a circulatory system or specialised gas exchange organs, because their feeding strategy involves one-way pumping of water through their porous bodies using flagellated collar cells. Each cell of the sponge's body is therefore exposed to a constant flow of fresh oxygenated water. They can therefore rely on diffusion across their cell membranes to carry out the gas exchange needed for respiration. In organisms that have circulatory systems associated with their specialized gas-exchange surfaces, a great variety of systems are used for the interaction between the two. In a countercurrent flow system, air (or, more usually, the water containing dissolved air) is drawn in the opposite direction to the flow of blood in the gas exchanger. A countercurrent system such as this maintains a steep concentration gradient along the length of the gas-exchange surface (see lower diagram in Fig. 2). This is the situation seen in the gills of fish and many other aquatic creatures. Although this theoretically allows almost complete transfer of a respiratory gas from one side of the exchanger to the other, in fish less than 80% of the oxygen in the water flowing over the gills is generally transferred to the blood. and dead-end air-filled sac systems found in the lungs of mammals. In a cocurrent flow system, the blood and gas (or the fluid containing the gas) move in the same direction through the gas exchanger. This means the magnitude of the gradient is variable along the length of the gas-exchange surface, and the exchange will eventually stop when an equilibrium has been reached (see upper diagram in Fig. 2). Cocurrent flow gas exchange systems are not known to be used in nature. ==Mammals==

The gas exchanger in mammals is internalized to form lungs, as it is in most of the larger land animals. Gas exchange occurs in microscopic dead-end air-filled sacs called alveoli, where a very thin membrane (called the blood-air barrier) separates the blood in the alveolar capillaries (in the walls of the alveoli) from the alveolar air in the sacs. and alveolar type I epithelial cells (or type 1 pneumocytes). The two red objects labeled "RBC" are red blood cells in the alveolar capillary blood. == Exchange membrane ==

The membrane across which gas exchange takes place in the alveoli (i.e. the blood-air barrier) is extremely thin (in humans, on average, 2.2 μm thick). The large surface area of the membrane comes from the folding of the membrane into about 300 million alveoli, with diameters of approximately 75–300 μm each. This provides an extremely large surface area (approximately 145 m2) across which gas exchange can occur. It is the first air to re-enter the alveoli during inhalation. Only after the dead space air has returned to the alveoli does the remainder of the tidal volume (500 ml - 150 ml = 350 ml) enter the alveoli. The composition of the air in the FRC is carefully monitored, by measuring the partial pressures of oxygen and carbon dioxide in the arterial blood. If either gas pressure deviates from normal, reflexes are elicited that change the rate and depth of breathing in such a way that normality is restored within seconds or minutes. compared to the concentration of oxygen in saturated arterial blood of about 9 mM (or 20 ml per 100 ml blood). thus blowing off too much CO2 from the blood into the outside air, precipitating a set of distressing symptoms which result from an excessively high pH of the extracellular fluids. Oxygen has a very low solubility in water, and is therefore carried in the blood loosely combined with hemoglobin. The oxygen is held on the hemoglobin by four ferrous iron-containing heme groups per hemoglobin molecule. When all the heme groups carry one O2 molecule each the blood is said to be "saturated" with oxygen, and no further increase in the partial pressure of oxygen will meaningfully increase the oxygen concentration of the blood. Most of the carbon dioxide in the blood is carried as HCO3− ions in the plasma. However the conversion of dissolved CO2 into HCO3− (through the addition of water) is too slow for the rate at which the blood circulates through the tissues on the one hand, and alveolar capillaries on the other. The reaction is therefore catalyzed by carbonic anhydrase, an enzyme inside the red blood cells. The reaction can go in either direction depending on the prevailing partial pressure of carbon dioxide. A small amount of carbon dioxide is carried on the protein portion of the hemoglobin molecules as carbamino groups. The total concentration of carbon dioxide (in the form of bicarbonate ions, dissolved CO2, and carbamino groups) in arterial blood (i.e. after it has equilibrated with the alveolar air) is about 26 mM (or 58 ml/100 ml), compared to the concentration of oxygen in saturated arterial blood of about 9 mM (or 20 ml/100 ml blood). ==Other vertebrates==

Fish The dissolved oxygen content in fresh water is approximately 8–10 milliliters per liter compared to that of air which is 210 milliliters per liter. Water is 800 times more dense than air and 100 times more viscous. Gill rakers are found within the exchange system in order to filter out food, and keep the gills clean. Gills use a countercurrent flow system that increases the efficiency of oxygen-uptake (and waste gas loss). The lower floor of the mouth is moved in a "pumping" manner, which can be observed by the naked eye. Reptiles All reptiles breathe using lungs. In squamates (the lizards and snakes) ventilation is driven by the axial musculature, but this musculature is also used during movement, so some squamates rely on buccal pumping to maintain gas exchange efficiency. Due to the rigidity of turtle and tortoise shells, significant expansion and contraction of the chest is difficult. Turtles and tortoises depend on muscle layers attached to their shells, which wrap around their lungs to fill and empty them. Some aquatic turtles can also pump water into a highly vascularised mouth or cloaca to achieve gas-exchange. Crocodiles have a structure similar to the mammalian diaphragm - the diaphragmaticus - but this muscle helps create a unidirectional flow of air through the lungs rather than a tidal flow: this is more similar to the air-flow seen in birds than that seen in mammals. During inhalation, the diaphragmaticus pulls the liver back, inflating the lungs into the space this creates. Air flows into the lungs from the bronchus during inhalation, but during exhalation, air flows out of the lungs into the bronchus by a different route: this one-way movement of gas is achieved by aerodynamic valves in the airways. Birds Birds have lungs but no diaphragm. They rely mostly on air sacs for ventilation. These air sacs do not play a direct role in gas exchange, but help to move air unidirectionally across the gas exchange surfaces in the lungs. During inhalation, fresh air is taken from the trachea down into the posterior air sacs and into the parabronchi which lead from the posterior air sacs into the lung. The air that enters the lungs joins the air which is already in the lungs, and is drawn forward across the gas exchanger into anterior air sacs. During exhalation, the posterior air sacs force air into the same parabronchi of the lungs, flowing in the same direction as during inhalation, allowing continuous gas exchange irrespective of the breathing cycle. Air exiting the lungs during exhalation joins the air being expelled from the anterior air sacs (both consisting of "spent air" that has passed through the gas exchanger) entering the trachea to be exhaled (Fig. 10). Selective bronchoconstriction at the various bronchial branch points ensures that the air does not ebb and flow through the bronchi during inhalation and exhalation, as it does in mammals. The unidirectional airflow through the parabronchi exchanges respiratory gases with a crosscurrent blood flow (Fig. 9). The partial pressure of O2 (P_{{\mathrm{O}}_2}) in the parabronchioles declines along their length as O2 diffuses into the blood. The capillaries leaving the exchanger near the entrance of airflow take up more O2 than capillaries leaving near the exit end of the parabronchi. When the contents of all capillaries mix, the final P_{{\mathrm{O}}_2} of the mixed pulmonary venous blood is higher than that of the exhaled air, but lower than that of the inhaled air. ==Plants==

Gas exchange in plants is dominated by the roles of carbon dioxide, oxygen and water vapor. is the only carbon source for autotrophic growth by photosynthesis, and when a plant is actively photosynthesising in the light, it will be taking up carbon dioxide, and losing water vapor and oxygen. At night, plants respire, and gas exchange partly reverses: water vapor is still lost (but to a smaller extent), but oxygen is now taken up and carbon dioxide released. plant leaf, showing the key plant organs involved in gas exchange Plant gas exchange occurs mostly through the leaves. Gas exchange between a leaf and the atmosphere occurs simultaneously through two pathways: 1) epidermal cells and cuticular waxes (usually referred as 'cuticle') which are always present at each leaf surface, and 2) stomata, which typically control the majority of the exchange. Gases enter into the photosynthetic tissue of the leaf through dissolution onto the moist surface of the palisade and spongy mesophyll cells. The spongy mesophyll cells are loosely packed, allowing for an increased surface area, and consequently an increased rate of gas-exchange. Uptake of carbon dioxide necessarily results in some loss of water vapor, because both molecules enter and leave by the same stomata, so plants experience a gas exchange dilemma: gaining enough without losing too much water. Therefore, water loss from other parts of the leaf is minimised by the waxy cuticle on the leaf's epidermis. The size of a stoma is regulated by the opening and closing of its two guard cells: the turgidity of these cells determines the state of the stomatal opening, and this itself is regulated by water stress. Plants showing crassulacean acid metabolism are drought-tolerant xerophytes and perform almost all their gas-exchange at night, because it is only during the night that these plants open their stomata. By opening the stomata only at night, the water vapor loss associated with carbon dioxide uptake is minimised. However, this comes at the cost of slow growth: the plant has to store the carbon dioxide in the form of malic acid for use during the day, and it cannot store unlimited amounts. Gas exchange measurements are important tools in plant science: this typically involves sealing the plant (or part of a plant) in a chamber and measuring changes in the concentration of carbon dioxide and water vapour with an infrared gas analyzer. If the environmental conditions (humidity, concentration, light and temperature) are fully controlled, the measurements of uptake and water release reveal important information about the assimilation and transpiration rates. The intercellular concentration reveals important information about the photosynthetic condition of the plants. Simpler methods can be used in specific circumstances: hydrogencarbonate indicator can be used to monitor the consumption of in a solution containing a single plant leaf at different levels of light intensity, and oxygen generation by the pondweed Elodea can be measured by simply collecting the gas in a submerged test-tube containing a small piece of the plant. ==Invertebrates==

The mechanism of gas exchange in invertebrates depends their size, feeding strategy, and habitat (aquatic or terrestrial). , red: choanocytes, grey: mesohyl, pale blue: water flow The sponges (Porifera) are sessile creatures, meaning they are unable to move on their own and normally remain attached to their substrate. They obtain nutrients through the flow of water across their cells, and they exchange gases by simple diffusion across their cell membranes. Pores called ostia draw water into the sponge and the water is subsequently circulated through the sponge by cells called choanocytes which have hair-like structures that move the water through the sponge. The cnidarians include corals, sea anemones, jellyfish and hydras. These animals are always found in aquatic environments, ranging from fresh water to salt water. They do not have any dedicated respiratory organs; instead, every cell in their body can absorb oxygen from the surrounding water, and release waste gases to it. One key disadvantage of this feature is that cnidarians can die in environments where water is stagnant, as they deplete the water of its oxygen supply. Corals often form symbiosis with other organisms, particularly photosynthetic dinoflagellates. In this symbiosis, the coral provides shelter and the other organism provides nutrients to the coral, including oxygen. The roundworms (Nematoda), flatworms (Platyhelminthes), and many other small invertebrate animals living in aquatic or otherwise wet habitats do not have a dedicated gas-exchange surface or circulatory system. They instead rely on diffusion of and directly across their cuticle. The cuticle is the semi-permeable outermost layer of their bodies. Other aquatic invertebrates such as most molluscs (Mollusca) and larger crustaceans (Crustacea) such as lobsters, have gills analogous to those of fish, which operate in a similar way. Unlike the invertebrates groups mentioned so far, insects are usually terrestrial, and exchange gases across a moist surface in direct contact with the atmosphere, rather than in contact with surrounding water. The insect's exoskeleton is impermeable to gases, including water vapor, so they have a more specialised gas exchange system, requiring gases to be directly transported to the tissues via a complex network of tubes. This respiratory system is separated from their circulatory system. Gases enter and leave the body through openings called spiracles, located laterally along the thorax and abdomen. Similar to plants, insects are able to control the opening and closing of these spiracles, but instead of relying on turgor pressure, they rely on muscle contractions. These contractions result in an insect's abdomen being pumped in and out. The spiracles are connected to tubes called tracheae, which branch repeatedly and ramify into the insect's body. These branches terminate in specialised tracheole cells which provides a thin, moist surface for efficient gas exchange, directly with cells. The other main group of terrestrial arthropod, the arachnids (spiders, scorpion, mites, and their relatives) typically perform gas exchange with a book lung. ==Summary of main gas exchange systems==