Diagram of the human lungs with the respiratory tract visible, and different colours for each lobe
The human lungs flank the heart and great vessels in the chest cavity
The lungs are the primary organs of the respiratory system in humans and many other animals including a few fish and some snails. In mammals and most other vertebrates, two lungs are located near the backbone on either side of the heart. Their function in the respiratory system is to extract oxygen from the atmosphere and transfer it into the bloodstream, and to release carbon dioxide from the bloodstream into the atmosphere, in a process of gas exchange. Respiration is driven by different muscular systems in different species. Mammals, reptiles and birds use their different muscles to support and foster breathing. In early tetrapods, air was driven into the lungs by the pharyngeal muscles via buccal pumping, a mechanism still seen in amphibians. In humans, the main muscle of respiration that drives breathing is the diaphragm. The lungs also provide airflow that makes vocal sounds including human speech possible.
Humans have two lungs, a right lung and a left lung. They are situated within the thoracic cavity of the chest. The right lung is bigger than the left, which shares space in the chest with the heart. The lungs together weigh approximately 1.3 kilograms (2.9 lb), and the right is heavier. The lungs are part of the lower respiratory tract that begins at the trachea and branches into the bronchi and bronchioles, and which receive air breathed in via the conducting zone. The conducting zone ends at the terminal bronchioles. These divide into the respiratory bronchioles of the respiratory zone which divide into alveolar ducts that give rise to the microscopic alveoli, where gas exchange takes place. Together, the lungs contain approximately 2,400 kilometres (1,500 mi) of airways and 300 to 500 million alveoli. Each lung is enclosed within a pleural sac which allows the inner and outer walls to slide over each other whilst breathing takes place, without much friction. This sac also divides each lung into sections called lobes. The right lung has three lobes and the left has two. The lobes are further divided into bronchopulmonary segments and lobules. The lungs have a unique blood supply, receiving deoxygenated blood from the heart in the pulmonary circulation for the purposes of receiving oxygen and releasing carbon dioxide, and a separate supply of oxygenated blood to the tissue of the lungs, in the bronchial circulation.
The tissue of the lungs can be affected by a number of diseases, including pneumonia and lung cancer. Chronic obstructive pulmonary disease includes chronic bronchitis and previously termed emphysema, can be related to smoking or exposure to harmful substances such as coal dust, asbestos fibres and crystalline silica dust. Diseases such as bronchitis can also affect the respiratory tract. Medical terms related to the lung often begin with pulmo-, from the Latin pulmonarius (of the lungs) as in pulmonology, or with pneumo- (from Greek πνεύμων "lung") as in pneumonia.
In embryonic development, the lungs begin to develop as an outpouching of the foregut, a tube which goes on to form the upper part of the digestive system. When the lungs are formed the fetus is held in the fluid-filled amniotic sac and so they do not function to breathe. Blood is also diverted from the lungs through the ductus arteriosus. At birth however, air begins to pass through the lungs, and the diversionary duct closes, so that the lungs can begin to respire. The lungs only fully develop in early childhood.
- 1 Structure of the human lungs
- 2 Development
- 3 Function
- 4 Clinical significance
- 5 Other animals
- 6 Evolutionary origins
- 7 Foot note
- 8 See also
- 9 Further reading
- 10 References
Structure of the human lungs
The lungs are located in the chest on either side of the heart in the rib cage. They are conical in shape with a narrow rounded apex at the top, and a broad concave base that rests on the convex surface of the diaphragm. The apex of the lung extends into the root of the neck, reaching shortly above the level of the sternal end of the first rib. The lungs stretch from close to the backbone in the rib cage to the front of the chest and downwards from the lower part of the trachea to the diaphragm. The left lung shares space with the heart, and has an indentation in its border called the cardiac notch of the left lung to accommodate this. The front and outer sides of the lung face the ribs, which make light indentations on their surfaces. The medial surfaces of the lungs face towards the centre of the chest, and lie against the heart, great vessels, and the carina where the trachea divides into the two main bronchi. The cardiac impression is an indentation formed on the surfaces of the lungs where they rest against the heart.
The lungs are surrounded by the pulmonary pleurae. The pleurae are two serous membranes; the outer parietal pleura lines the inner wall of the rib cage and the inner visceral pleura directly lines the surface of the lungs. Between the pleurae is a potential space called the pleural cavity containing a thin layer of lubricating pleural fluid. Each lung is divided into lobes by the invaginations of the pleura as fissures. The fissures are double folds of pleura that section the lungs and help in their expansion.
The lobes of the lungs are further divided into bronchopulmonary segments based on the locations of bronchioles. Segments for the left and right lung are shown in the table. The segmental anatomy is useful clinically for localising disease processes in the lungs.
|Right lung||Left lung|
The right lung has both more lobes and segments than the left. It is divided into three lobes, an upper, middle, and a lower, by two fissures, one oblique and one horizontal. The upper, horizontal fissure, separates the upper from the middle lobe. It begins in the lower oblique fissure near the posterior border of the lung, and, running horizontally forward, cuts the anterior border on a level with the sternal end of the fourth costal cartilage; on the mediastinal surface it may be traced backward to the hilum.
The mediastinal surface of the right lung is indented by a number of nearby structures. The heart sits in an impression called the cardiac impression. Above the hilum of the lung is an arched groove for the azygos vein, and above this is a wide groove for the superior vena cava and right brachiocephalic vein; behind this, and close to the top of the lung is a groove for the brachiocephalic artery. There is a groove for the esophagus behind the hilum and the pulmonary ligament, and near the lower part of the esophageal groove is a deeper groove for the inferior vena cava before it enters the heart.
The left lung is divided into two lobes, an upper and a lower, by the oblique fissure, which extends from the costal to the mediastinal surface of the lung both above and below the hilum. The left lung, unlike the right, does not have a middle lobe, though it does have a homologous feature, a projection of the upper lobe termed the “lingula”. Its name means “little tongue”. The lingula on the left serves as an anatomic parallel to the right middle lobe, with both areas being predisposed to similar infections and anatomic complications. There are two bronchopulmonary segments of the lingula: superior and inferior.
The mediastinal surface of the left lung has a large cardiac impression where the heart sits. This is deeper and larger than that on the right lung, at which level the heart projects to the left.
On the same surface, immediately above the hilum, is a well-marked curved groove for the aortic arch, and a groove below it for the descending aorta. The left subclavian artery, a branch off the aortic arch, sits in a groove from the arch to near the apex of the lung. A shallower groove in front of the artery and near the edge of the lung, lodges the left brachiocephalic vein. The esophagus may sit in a wider shallow impression at the base of the lung.
The lower respiratory tract including the trachea, bronchi, and bronchioles are lined with respiratory epithelium. This is a ciliated epithelium interspersed with goblet cells which produce mucus, and club cells with actions similar to macrophages. Incomplete rings of cartilage in the trachea and smaller plates of cartilage in the bronchi, keep these airways open. Bronchioles are too narrow to support cartilage and their walls are of smooth muscle, and this is largely absent in the narrower respiratory bronchioles which are mainly just of epithelium. The respiratory tract ends in lobules. Each lobule consists of a respiratory bronchiole, which branches into alveolar ducts and alveolar sacs, which in turn divide into alveoli.
The epithelial cells throughout the respiratory tract secrete epithelial lining fluid (ELF), the composition of which is tightly regulated and determines how well mucociliary clearance works.:Section 4 pages 7–8 (Page 4–7ff)
Alveoli consist of two types of alveolar cell and an alveolar macrophage. The two types of cell are known as type I and type II alveolar cells (also known as pneumocytes). Types I and II make up the walls and alveolar septa. Type I cells provide 95% of the surface area of each alveoli and are flat ("squamous"), and Type II cells generally cluster in the corners of the alveoli and have a cuboidal shape. Despite this, cells occur in a roughly equal ratio of 1:1 or 6:4.
Type I are squamous epithelial cells that make up the alveolar wall structure. They have extremely thin walls that enable an easy gas exchange. These type I cells also make up the alveolar septa which separate each alveolus. The septa consist of an epithelial lining and associated basement membranes. Type I cells are not able to divide, and consequently rely on differentiation from Type II cells.
The lower respiratory tract is part of the respiratory system, and consists of the trachea and the structures below this including the lungs. The trachea receives air from the pharynx and travels down to a place where it splits (the carina) into a right and left bronchus. These supply air to the right and left lungs, splitting progressively into the secondary and tertiary bronchi for the lobes of the lungs, and into smaller and smaller bronchioles until they become the respiratory bronchioles. These in turn supply air through alveolar ducts into the alveoli, where the exchange of gases take place. Oxygen breathed in, diffuses through the walls of the alveoli into the enveloping capillaries and into the circulation, and carbon dioxide diffuses into the lungs to be breathed out.
The bronchi in the conducting zone are reinforced with hyaline cartilage in order to hold open the airways. The bronchioles have no cartilage and are surrounded instead by smooth muscle. Air is warmed to 37 °C (99 °F), humidified and cleansed by the conducting zone; particles from the air being removed by the cilia on the respiratory epithelium lining the passageways.
The lungs have a dual blood supply provided by a bronchial and a pulmonary circulation. The bronchial circulation supplies oxygenated blood to the airways of the lungs, through the bronchial arteries that leave the aorta. There are usually three arteries, two to the left lung and one to the right, and they branch alongside the bronchi and bronchioles. The pulmonary circulation carries deoxygenated blood from the heart to the lungs and returns the oxygenated blood to the heart to supply the rest of the body.
The blood volume of the lungs, is about 450 millilitres on average, about 9 per cent of the total blood volume of the entire circulatory system. This quantity can easily fluctuate from between one-half and twice the normal volume.
The lungs are supplied by nerves of the autonomic nervous system. Input from the parasympathetic nervous system occurs via the vagus nerve. When stimulated by acetylcholine, this causes constriction of the smooth muscle lining the bronchus and bronchioles, and increases the secretions from glands. The lungs also have a sympathetic tone from norepinephrine acting on the beta 2 receptors in the respiratory tract, which causes bronchodilation.
The larynx, trachea, bronchi and lungs that make up the respiratory tract, begin to form during the fourth week of embryogenesis from the lung bud which appears ventrally to the caudal portion of the foregut.
The respiratory tract has a branching structure like that of a tree. In the embryo this structure is developed in the process of branching morphogenesis, and is generated by the repeated splitting of the tip of the branch. In the development of the lungs (as in some other organs) the epithelium forms branching tubes.The lung has a left-right symmetry and each bud known as a bronchial bud grows out as a tubular epithelium that becomes a bronchus. Each bronchus branches into bronchioles. The branching is a result of the tip of each tube bifurcating. The branching process forms the bronchi, bronchioles, and ultimately the alveoli. The four genes mostly associated with branching morphogenesis in the lung are the intercellular signalling protein – sonic hedgehog (SHH), fibroblast growth factors FGF10 and FGFR2b, and bone morphogenetic protein BMP4. FGF10 is seen to have the most prominent role. FGF10 is a paracrine signalling molecule needed for epithelial branching, and SHH inhibits FGF10. The development of the alveoli is influenced by a different mechanism whereby continued bifurcation is stopped and the distal tips become dilated to form the alveoli.
At the end of the fourth week the lung bud divides into two, the right and left primary bronchial buds on each side of the trachea. During the fifth week the right bud branches into three secondary bronchial buds and the left branches into two secondary bronchial buds. These give rise to the lobes of the lungs, three on the right and two on the left. Over the following week, the secondary buds branch into tertiary buds, about ten on each side. From the sixth week to the sixteenth week, the major elements of the lungs appear except the alveoli. From week 16 to week 26, the bronchi enlarge and lung tissue becomes highly vascularised. Bronchioles and alveolar ducts also develop. By week 26 the terminal bronchioles have formed which branch into two respiratory bronchioles. During the period covering the 26th week until birth the important blood–air barrier is established. Specialised type I alveolar cells where gas exchange will take place, together with the type II alveolar cells that secrete pulmonary surfactant, appear. The surfactant reduces the surface tension at the air-alveolar surface which allows expansion of the alveolar sacs. The alveolar sacs contain the primitive alveoli that form at the end of the alveolar ducts, and their appearance around the seventh month marks the point at which limited respiration would be possible, and the premature baby could survive.
At birth, the baby's lungs are filled with fluid secreted by the lungs and are not inflated. After birth the infant's central nervous system reacts to the sudden change in temperature and environment. This triggers the first breath, within about 10 seconds after delivery. Before birth, the lungs are filled with fetal lung fluid. After the first breath, the fluid is quickly absorbed into the body or exhaled. The resistance in the lung's blood vessels decreases giving an increased surface area for gas exchange, and the lungs begins to breathe spontaneously. This accompanies other changes which result in an increased amount of blood entering the lung tissues.
At birth the lungs are very undeveloped with only around one sixth of the alveoli of the adult lung present. The alveoli continue to form into early adulthood, and their ability to form when necessary is seen in the regeneration of the lung. Alveolar septa have a double capillary network instead of the single network of the developed lung. Only after the maturation of the capillary network can the lung enter a normal phase of growth. Following the early growth in numbers of alveoli there is another stage of the alveoli being enlarged.
The lungs are not capable of expanding themselves, and will only do so when there is an increase in the volume of the thoracic cavity. This is achieved by the muscles of respiration, through the contraction of the diaphragm, and the intercostal muscles which pull the rib cage upwards as shown in the diagram. During breathing out, at rest, the muscles of inhalation relax, returning the chest and abdomen to a resting position determined by their anatomical elasticity. At this point the lungs contain the functional residual capacity (FRC) of air, which, in the adult human, has a volume of about 2.5–3.0 litres.
During heavy breathing as, for instance, during exercise, a large number of accessory muscles in the neck and abdomen are recruited. Exhalation, instead of being passive, is now caused by the rib cage being actively pulled downwards at the front and sides by the abdominal muscles. This has the effect of decreasing the size of the rib cage, and of pushing the abdominal organs up against the diaphragm which bulges into the thorax. The lung volume of air at the end of exhalation is now less than the resting FRC. However, the lungs cannot be emptied completely and in an adult human there is always still at least one litre of residual air left in the lungs after maximum exhalation. The FRC always remains in the alveoli after a normal out-breath. With each breath only about 350 ml (i.e. less than 15%) of this alveolar air is expelled into the ambient air (with about 150 ml remaining behind in the airways, as "dead space" ventilation, as it is the first air to re-enter the alveoli on inhalation). This is immediately replaced with the same volume of fresh, but moistened, atmospheric air. It is therefore obvious that the 350 ml inhaled fresh air is highly diluted by the 3 litres of functional residual capacity air (i.e. the air that remains in the lungs after a normal exhalation), and that the composition of the alveolar air therefore changes very little under normal circumstances: the alveolar partial pressure of oxygen remains very close to 14 kPa ( 105 mmHg ), and that of carbon dioxide varies minimally around 5.3 kPa ( 40 mmHg ) throughout the respiratory cycle (of inhalation and exhalation).
The major function of the lungs is gas exchange between the blood gases and the alveolar air. The alveolar and pulmonary capillary gases equilibrate across the blood–air barrier, a very thin diffusion membrane which is only, on average, about 2 μm thick, consisting of the walls of the pulmonary alveoli, consisting of the alveolar epithelial cells, their basement membranes and the endothelial cells of the pulmonary capillaries. This membrane is folded into about 300 million small air sacs called alveoli (each between 75 and 300 µm in diameter) branching off from the bronchioles in the lungs, thus providing an extremely large surface area (estimates varying between 70 and 145 m2) for gas exchange to occur.
Since the blood gasses in the alveolar capillaries equilibrate with those in the alveolar air, the arterial blood that is spread evenly throughout the body by the left ventricle, will have the same partial pressures of oxygen and carbon dioxide as persist in the alveoli.
Control of breathing
The arterial partial pressures of oxygen and carbon dioxide in the arterial blood are homeostatically controlled. A rise in the arterial partial pressure of carbon dioxide, and, to a lesser extent, a fall in the arterial partial pressure of oxygen, will reflexly cause deeper and faster breathing till the blood gas tensions return to normal. The converse happens when the carbon dioxide tension falls, or, again to a lesser extent, the oxygen tension rises: the rate and depth of breathing are reduced till blood gas normality is restored.
It is only as a result of accurately maintaining the composition of the 3 litres alveolar air that, with each breath, some carbon dioxide is discharged into the atmosphere and some oxygen is taken up from the outside air. If more carbon dioxide than usual has been lost by a short period of hyperventilation, breathing will be slowed down or halted until the alveolar partial pressure of carbon dioxide has returned to 5.3 kPa ( 40 mmHg ). The opposite occurs after breath-holding.
The partial pressures of oxygen and carbon dioxide in the arterial blood are measured by the peripheral chemoreceptors – the aortic and carotid bodies, and the central chemoreceptors of the medulla oblongata of the brainstem. The peripheral chemoreceptors are more sensitive to the arterial partial pressure of oxygen than they are to the arterial partial pressure of carbon dioxide. The central chemoreceptor is particularly sensitive to the pH of the cerebrospinal fluid, which is directly influenced by the partial pressure of carbon dioxide in the arterial blood.
Information from the peripheral chemoreceptors is relayed to a series of interconnected nuclei which comprise the respiratory centres in the medulla oblongata and the pons of the brainstem. This information determines the average rate of ventilation of the alveoli of the lungs, to keep the partial pressures of oxygen and carbon dioxide in the arterial blood constant. The respiratory centre does so via motor neurons which activate the muscles of respiration (in particular the diaphragm).
Exercise also increases the respiratory rate, partly in response to the movement of the limbs detected by proprioceptors in the muscles and joints, an increase in body temperature, the release of adrenaline (epinephrine) from the adrenal glands, and from motor impulses originating from the cerebral cortex.
The lungs possess several characteristics which protect against infection. The lung tract is lined by epithelia with hair-like projections called cilia that beat rhythmically and carry mucus. This mucociliary clearance is an important defence system against air-borne infection. The dust particles and bacteria in the inhaled air are caught in the mucosal surface of respiratory passages and are moved up towards the pharynx by the rhythmic upward beating action of the cilia. The lining of the lung also secretes immunoglobulin A which protects against respiratory infections; goblet cells secrete mucus which also contains several antimicrobial compounds such as defensins, antiproteases, and antioxidates. In addition, the lining of the lung also contains macrophages, immune cells which engulf and destroy debris and microbes that enter the lung in a process known as phagocytosis; and dendritic cells which present antigens to activate components of the adaptive immune system such as T-cells and B-cells.
The size of the respiratory tract and the flow of air also protect the lungs from larger particles. Smaller particles deposit in the mouth and behind the mouth in the oropharynx, and larger particles are trapped in nasal hair after inhalation.
In addition to their function in respiration, the lungs also have a number of other functions. They are involved in maintaining homeostasis, helping in the regulation of blood pressure as part of the renin-angiotensin system. The inner lining of the blood vessels secretes angiotensin-converting enzyme (ACE) an enzyme that catalyses the conversion of angiotensin I to angiotensin II. The lungs are involved in the blood's acid-base homeostasis by expelling carbon dioxide when breathing.
The lungs also serve a protective role. Several blood-borne substances, such as a few types of prostaglandins, leukotrienes, serotonin and bradykinin, are excreted through the lungs. Drugs and other substances can be absorbed, modified or excreted in the lungs. The lungs filter out small blood clots from veins and prevent them from entering arteries and causing strokes.
The lungs also play a pivotal role in speech by providing air and airflow for the creation of vocal sounds. Lungs also provide the airflow that enables the expression of many emotions such as sighing, yawning, sobbing, laughing in humans and the vocal sounds in other animals.
Lungs can be affected by a variety of diseases. Pulmonology is the medical speciality that deals with diseases involving the respiratory tract, and cardiothoracic surgery is the surgical field that deals with surgery of the lungs.
Inflammatory conditions of the lung tissue are pneumonia, of the respiratory tract are bronchitis and bronchiolitis, and of the pleurae surrounding the lungs pleurisy. Inflammation is usually caused by infections due to bacteria or viruses. When the lung tissue is inflamed due to other causes it is called pneumonitis. One major cause of bacterial pneumonia is tuberculosis. Chronic infections often occur in those with immunodeficiency and can include a fungal infection by Aspergillus fumigatus that can lead to an aspergilloma forming in the lung.
A pulmonary embolism is a blood clot that becomes lodged in the pulmonary arteries. The majority of emboli arise because of deep vein thrombosis in the legs. Pulmonary emboli may be investigated using a ventilation/perfusion scan, a CT scan of the arteries of the lung, or blood tests such as the D-dimer. Pulmonary hypertension describes an increased pressure at the beginning of the pulmonary artery that has a large number of differing causes. Other rarer conditions may also affect the blood supply of the lung, such as granulomatosis with polyangiitis, which causes inflammation of the small blood vessels of the lungs and kidneys.
A lung contusion is a bruise caused by chest trauma. It results in hemorrhage of the alveoli causing a build-up of fluid which can impair breathing, and this can be either mild or severe. The function of the lungs can also be affected by compression from fluid in the pleural cavity pleural effusion, or other substances such as air (pneumothorax), blood (hemothorax), or rarer causes. These may be investigated using a chest X-ray or CT scan, and may require the insertion of a surgical drain until the underlying cause is identified and treated.
Asthma, chronic bronchitis, bronchiectasis and chronic obstructive pulmonary disease (COPD) are all obstructive lung diseases characterised by airway obstruction. This limits the amount of air that is able to enter alveoli because of constriction of the bronchial tree, due to inflammation. Obstructive lung diseases are often identified because of symptoms and diagnosed with pulmonary function tests such as spirometry. Many obstructive lung diseases are managed by avoiding triggers (such as dust mites or smoking), with symptom control such as bronchodilators, and with suppression of inflammation (such as through corticosteroids) in severe cases. One common cause of COPD and emphysema is smoking, and common causes of bronchiectasis include severe infections and cystic fibrosis. The definitive cause of asthma is not yet known.
Some types of chronic lung diseases are classified as restrictive lung disease, because of a restriction in the amount of lung tissue involved in respiration. These include pulmonary fibrosis which can occur when the lung is inflamed for a long period of time. Fibrosis in the lung replaces functioning lung tissue with fibrous connective tissue. This can be due to a large variety of occupational diseases such as Coalworker's pneumoconiosis, autoimmune diseases or more rarely to a reaction to medication.
Lung cancer can either arise directly from lung tissue or as a result of metastasis from another part of the body. There are two main types of primary tumour described as either small-cell or non-small-cell lung carcinomas. The major risk factor for cancer is smoking. Once a cancer is identified it is staged using scans such as a CT scan and a sample of tissue (a biopsy) is taken. Cancers may be treated by surgically removing the tumour, radiotherapy, chemotherapy or combinations thereof, or with the aim of symptom control.Lung cancer screening is being recommended in the United States for high-risk populations.
Congenital disorders include cystic fibrosis, pulmonary hypoplasia (an incomplete development of the lungs)congenital diaphragmatic hernia, and infant respiratory distress syndrome caused by a deficiency in lung surfactant. An azygos lobe is a congenital anatomical variation which though usually without effect can cause problems in thoracoscopic procedures.
A pneumothorax (collapsed lung) is an abnormal collection of air in the pleural space that causes an uncoupling of the lung from the chest wall. The lung cannot expand against the air pressure inside the pleural space. An easy to understand example is a traumatic pneumothorax, where air enters the pleural space from outside the body, as occurs with puncture to the chest wall. Similarly, a scuba diver ascending while holding their breath with their lungs fully inflated can cause air sacs (alveoli) to burst and leak high pressure air into the pleural space.
Lung function testing
Lung function testing is carried out by evaluating a person's capacity to inhale and exhale in different circumstances. The volume of air inhaled and exhaled by a person at rest is the tidal volume (normally 500-750mL); the inspiratory reserve volume and expiratory reserve volume are the additional amounts a person is able to forcibly inhale and exhale respectively. The summed total of forced inspiration and expiration is a person's vital capacity. Not all air is expelled from the lungs even after a forced breath out; the remainder of the air is called the residual volume. Together these terms are referred to as lung volumes.
Pulmonary plethysmographs are used to measure functional residual capacity. Functional residual capacity cannot be measured by tests that rely on breathing out, as a person is only able to breathe a maximum of 80% of their total functional capacity. The total lung capacity depends on the person's age, height, weight, and sex, and normally ranges between 4 and 6 litres. Females tend to have a 20–25% lower capacity than males. Tall people tend to have a larger total lung capacity than shorter people. Smokers have a lower capacity than nonsmokers. Thinner persons tend to have a larger capacity, and capacity can be increased by physical training as much as 40%.
Other lung function tests include spirometry, measuring the amount (volume) and flow of air that can be inhaled and exhaled. The maximum volume of breath that can be exhaled is called the vital capacity. In particular, how much a person is able to exhale in one second (called forced expiratory volume (FEV1) as a proportion of how much they are able to exhale in total (FEV). This ratio, the FEV1/FEV ratio, is important to distinguish whether a lung disease is restrictive or obstructive. Another test is that of the lung's diffusing capacity – this is a measure of the transfer of gas from air to the blood in the lung capillaries.
The lungs of birds are relatively small, but are connected to 8–9 air sacs that extend through much of the body, and are in turn connected to air spaces within the bones. On inhalation, air travels through the trachea of a bird into the 8–9 air sacs. Air then travels continuously from the air sacs at the back, through the lungs, which are relatively fixed in size, to the air sacs at the front. From here, the air is exhaled. These fixed size lungs are called "circulatory lungs", as distinct from the "bellows-type lungs" found in most other animals.
The lungs of birds contain millions of tiny parallel passages called parabronchi. Small sacs called atria radiate from the walls of the tiny passages, like the alveoli in other lungs, are the site of gas exchange by simple diffusion. The blood flow around the parabronchi and their atria, forms a cross-current process of gas exchange (see diagram on the right).
The air sacs, which hold air, do not contribute much to gas exchange, despite being thin-walled, as they are poorly vascularised. The air sacs expand and contract due to changes in the volume in the thorax and abdomen. This volume change is caused by the movement of the sternum and ribs and this movement is often synchronised with movement of the flight muscles.
Parabronchi in which the air flow is unidirectional are called paleopulmonic parabronchi and are found in all birds. Some birds, however, have, in addition, a lung structure where the air flow in the parabronchi is bidirectional. These are termed neopulmonic parabronchi.
The lungs of most reptiles have a single bronchus running down the centre, from which numerous branches reach out to individual pockets throughout the lungs. These pockets are similar to alveoli in mammals, but much larger and fewer in number. These give the lung a sponge-like texture. In tuataras, snakes, and some lizards, the lungs are simpler in structure, similar to that of typical amphibians.
Snakes and limbless lizards typically possess only the right lung as a major respiratory organ; the left lung is greatly reduced, or even absent. Amphisbaenians, however, have the opposite arrangement, with a major left lung, and a reduced or absent right lung.
Both crocodilians and monitor lizards have developed lungs similar to those of birds, providing an unidirectional airflow and even possessing air sacs. The now extinct pterosaurs have seemingly even further refined this type of lung, extending the airsacs into the wing membranes and, in the case of lonchodectids, tupuxuara, and azhdarchoids, the hindlimbs.
Reptilian lungs typically receive air via expansion and contraction of the ribs driven by axial muscles and buccal pumping. Crocodilians also rely on the hepatic piston method, in which the liver is pulled back by a muscle anchored to the pubic bone (part of the pelvis) called the diaphragmaticus, which in turn creates negative pressure in the crocodile's thoracic cavity, allowing air to be moved into the lungs by Boyle's law. Turtles, which are unable to move their ribs, instead use their forelimbs and pectoral girdle to force air in and out of the lungs.
The lungs of most frogs and other amphibians are simple and balloon-like, with gas exchange limited to the outer surface of the lung. This is not very efficient, but amphibians have low metabolic demands and can also quickly dispose of carbon dioxide by diffusion across their skin in water, and supplement their oxygen supply by the same method. Amphibians employ a positive pressure system to get air to their lungs, forcing air down into the lungs by buccal pumping. This is distinct from most higher vertebrates, who use a breathing system driven by negative pressure where the lungs are inflated by expanding the rib cage. In buccal pumping, the floor of the mouth is lowered, filling the mouth cavity with air. The throat muscles then presses the throat against the underside of the skull, forcing the air into the lungs.
Due to the possibility of respiration across the skin combined with small size, all known lungless tetrapods are amphibians. The majority of salamander species are lungless salamanders, which respirate through their skin and tissues lining their mouth. This necessarily restrict their size: all are small and rather thread-like in appearance, maximising skin surface relative to body volume. Other known lungless tetrapods are the Bornean flat-headed frog and Atretochoana eiselti, a caecilian.
The lungs of amphibians typically have a few narrow internal walls (septa) of soft tissue around the outer walls, increasing the respiratory surface area and giving the lung a honey-comb appearance. In some salamanders even these are lacking, and the lung has a smooth wall. In caecilians, as in snakes, only the right lung attains any size or development.
The lungs of lungfish are similar to those of amphibians, with few, if any, internal septa. In the Australian lungfish, there is only a single lung, albeit divided into two lobes. Other lungfish and Polypterus, however, have two lungs, which are located in the upper part of the body, with the connecting duct curving around and above the esophagus. The blood supply also twists around the esophagus, suggesting that the lungs originally evolved in the ventral part of the body, as in other vertebrates.
Some invertebrates have "lungs" that serve a similar respiratory purpose as, but are not evolutionarily related to, vertebrate lungs. Some arachnids, spiders and scorpions, have structures called "book lungs" used for atmospheric gas exchange. Some species of spider have four pairs of book lungs but most have two pairs. Scorpions have spiracles on their body for the entrance of air to the book lungs.
The coconut crab is terrestrial and uses structures called branchiostegal lungs to breathe air. They cannot swim and would drown in water, yet they possess a rudimentary set of gills. They can breathe on land and hold their breath underwater. The branchiostegal lungs are seen as a developmental adaptive stage from water-living to enable land-living, or from fish to amphibian.
Pulmonates are mostly land snails and slugs that have developed a simple lung from the mantle cavity. An externally located opening called the pneumostome allows air to be taken into the mantle cavity lung.
The lungs of today's terrestrial vertebrates and the gas bladders of today's fish are believed to have evolved from simple sacs, as outpocketings of the esophagus, that allowed early fish to gulp air under oxygen-poor conditions. These outpocketings first arose in the bony fish. In most of the ray-finned fish the sacs evolved into closed off gas bladders, while a number of carps, trouts, herrings, catfish, and eels have retained the physostome condition with the sack being open to the esophagus. In more basal bony fish, such as the gar, bichir, bowfin and the lobe-finned fish, the bladders have evolved to primarily function as lungs. The lobe-finned fish gave rise to the land-based tetrapods. Thus, the lungs of vertebrates are homologous to the gas bladders of fish (but not to their gills).
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