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The lungs are the primary organs of the
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies g ...
in
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s and most other
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
s, including some
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class ...
s and a small number of
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
. In
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s and most other
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, two lungs are located near the backbone on either side of the
heart The heart is a muscular Organ (biology), organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as ca ...
. Their function in the respiratory system is to extract
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...
from the air and transfer it into the bloodstream, and to release
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
from the bloodstream into the atmosphere, in a process of
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a ...
. Respiration is driven by different muscular systems in different species. Mammals,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
s and
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s use their different muscles to support and foster breathing. In earlier
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s, 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 Diaphragm may refer to: Anatomy * Thoracic diaphragm, a thin sheet of muscle between the thorax and the abdomen * Pelvic diaphragm or pelvic floor, a pelvic structure * Urogenital diaphragm or triangular ligament, a pelvic structure Other * Diap ...
. The lungs also provide airflow that makes vocal sounds including human speech possible. Humans have two lungs, one on the left and one on the right. They are situated within the thoracic cavity of the chest. The right lung is bigger and heavier than the left, which shares space in the chest with the heart. The lungs together weigh approximately . The lungs are part of the lower respiratory tract that begins at the
trachea The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air- breathing animals with lungs. The trachea extends from t ...
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 alveolar sacs that contain the
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * M ...
, where gas exchange takes place. Alveoli are also sparsely present on the walls of the respiratory bronchioles and alveolar ducts. Together, the lungs contain approximately of airways and 300 to 500 million alveoli. Each lung is enclosed within a pleural sac of two
membranes A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. Bi ...
called pleurae; the membranes are separated by a film of pleural fluid, which allows the inner and outer membranes to slide over each other whilst breathing takes place, without much friction. The inner pleura 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 segment A bronchopulmonary segment is a portion of lung supplied by a specific segmental bronchus and its vessels. These arteries branch from the pulmonary and bronchial arteries, and run together through the center of the segment. Veins and lymphatic v ...
s and pulmonary 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 respiratory diseases, including
pneumonia Pneumonia is an inflammatory condition of the lung primarily affecting the small air sacs known as alveoli. Symptoms typically include some combination of productive or dry cough, chest pain, fever, and difficulty breathing. The severit ...
and
lung cancer Lung cancer, also known as lung carcinoma (since about 98–99% of all lung cancers are carcinomas), is a malignant lung tumor characterized by uncontrolled cell growth in tissues of the lung. Lung carcinomas derive from transformed, mali ...
.
Chronic obstructive pulmonary disease Chronic obstructive pulmonary disease (COPD) is a type of progressive lung disease characterized by long-term respiratory symptoms and airflow limitation. The main symptoms include shortness of breath and a cough, which may or may not produce ...
includes chronic bronchitis and
emphysema Emphysema, or pulmonary emphysema, is a lower respiratory tract disease, characterised by air-filled spaces ( pneumatoses) in the lungs, that can vary in size and may be very large. The spaces are caused by the breakdown of the walls of the alv ...
, and can be related to
smoking Smoking is a practice in which a substance is burned and the resulting smoke is typically breathed in to be tasted and absorbed into the bloodstream. Most commonly, the substance used is the dried leaves of the tobacco plant, which have b ...
or exposure to harmful substances. A number of occupational lung diseases can be caused by substances such as coal dust, asbestos fibres, and
crystal A crystal or crystalline solid is a solid material whose constituents (such as atoms, molecules, or ions) are arranged in a highly ordered microscopic structure, forming a crystal lattice that extends in all directions. In addition, macro ...
line
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is ...
dust. Diseases such as bronchitis can also affect the respiratory tract. Medical terms related to the lung often begin with ''pulmo-'', from the
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''pulmonarius'' (of the lungs) as in pulmonology, or with ''pneumo-'' (from Greek πνεύμων "lung") as in
pneumonia Pneumonia is an inflammatory condition of the lung primarily affecting the small air sacs known as alveoli. Symptoms typically include some combination of productive or dry cough, chest pain, fever, and difficulty breathing. The severit ...
. 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 A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal dev ...
is held in the fluid-filled
amniotic sac The amniotic sac, also called the bag of waters or the membranes, is the sac in which the embryo and later fetus develops in amniotes. It is a thin but tough transparent pair of membranes that hold a developing embryo (and later fetus) until sh ...
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.


Structure


Anatomy

The lungs are located in the chest on either side of the
heart The heart is a muscular Organ (biology), organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as ca ...
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 Diaphragm may refer to: Anatomy * Thoracic diaphragm, a thin sheet of muscle between the thorax and the abdomen * Pelvic diaphragm or pelvic floor, a pelvic structure * Urogenital diaphragm or triangular ligament, a pelvic structure Other * Diap ...
. 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 rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a ...
. 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 lungs 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. Both lungs have a central recession called the hilum, where the
blood vessel The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide awa ...
s and airways pass into the lungs making up the root of the lung. There are also bronchopulmonary lymph nodes on the hilum. 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.


Lobes

Each lung is divided into sections called lobes by the infoldings of the visceral pleura as fissures. Lobes are divided into segments, and segments have further divisions as lobules. There are three lobes in the right lung and two lobes in the left lung.


Fissures

The fissures are formed in early prenatal development by invaginations of the visceral pleura that divide the lobar bronchi, and section the lungs into lobes that helps in their expansion. The right lung is divided into three lobes by a horizontal fissure, and an oblique fissure. The left lung is divided into two lobes by an oblique fissure which is closely aligned with the oblique fissure in the right lung. In the right lung the upper horizontal fissure, separates the upper (superior) lobe from the middle lobe. The lower, oblique fissure separates the lower lobe from the middle and upper lobes. Variations in the fissures are fairly common being either incompletely formed or present as an extra fissure as in the azygos fissure, or absent. Incomplete fissures are responsible for interlobar collateral ventilation, airflow between lobes which is unwanted in some
lung volume reduction Emphysema, or pulmonary emphysema, is a lower respiratory tract disease, characterised by air-filled spaces ( pneumatoses) in the lungs, that can vary in size and may be very large. The spaces are caused by the breakdown of the walls of the alv ...
procedures.


Segments

The main or primary bronchi enter the lungs at the hilum and initially branch into secondary bronchi also known as lobar bronchi that supply air to each lobe of the lung. The lobar bronchi branch into tertiary bronchi also known as segmental bronchi and these supply air to the further divisions of the lobes known as
bronchopulmonary segment A bronchopulmonary segment is a portion of lung supplied by a specific segmental bronchus and its vessels. These arteries branch from the pulmonary and bronchial arteries, and run together through the center of the segment. Veins and lymphatic v ...
s. Each bronchopulmonary segment has its own (segmental) bronchus and arterial supply. 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. A segment is a discrete unit that can be surgically removed without seriously affecting surrounding tissue.


Right 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 lobe 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 The mediastinum (from ) is the central compartment of the thoracic cavity. Surrounded by loose connective tissue, it is an undelineated region that contains a group of structures within the thorax, namely the heart and its vessels, the esopha ...
surface it may be traced back to the hilum. The lower, oblique fissure, separates the lower from the middle and upper lobes and is closely aligned with the oblique fissure in the left lung. 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 The left and right brachiocephalic veins (previously called innominate veins) are major veins in the upper chest, formed by the union of each corresponding internal jugular vein and subclavian vein. This is at the level of the sternoclavicular ...
; 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.Alt URL
/ref> The weight of the right lung varies between individuals, with a standard reference range in men of and in women of .


Left lung

The left lung is divided into two lobes, an upper and a lower lobe, 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 lung serves as an anatomic parallel to the middle lobe on the right lung, with both areas being predisposed to similar infections and anatomic complications. There are two
bronchopulmonary segment A bronchopulmonary segment is a portion of lung supplied by a specific segmental bronchus and its vessels. These arteries branch from the pulmonary and bronchial arteries, and run together through the center of the segment. Veins and lymphatic v ...
s 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 The aortic arch, arch of the aorta, or transverse aortic arch () is the part of the aorta between the ascending and descending aorta. The arch travels backward, so that it ultimately runs to the left of the trachea. Structure The aorta begins ...
, 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 left and right brachiocephalic veins (previously called innominate veins) are major veins in the upper chest, formed by the union of each corresponding internal jugular vein and subclavian vein. This is at the level of the sternoclavicular ...
. The
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to t ...
may sit in a wider shallow impression at the base of the lung. The weight of the left lung, by standard reference range, in men is in women .


Illustrations

File:CT-Thorax-5.0-B70f-Lungs.jpg, Chest CT (axial lung window) File:COR-2-STND-CHEST-LUNGS.jpg, Chest CT (coronal lung window) File:CT-Thorax-5.0-B70f.ogg, Chest CT (axial lung window) File:COR-2-STND-CHEST.ogg, Chest CT (coronal lung window) File:Meet the lungs.webm, "Meet the lungs" from
Khan Academy Khan Academy is an American non-profit educational organization created in 2008 by Sal Khan. Its goal is creating a set of online tools that help educate students. The organization produces short lessons in the form of videos. Its website also i ...
File:MP1 Pulmonology.webm, Pulmonology Video File:Lobes of the Lung ogg.mov (1).ogg, 3D anatomy of the lung lobes and fissures.


Microanatomy

The lungs are part of the lower respiratory tract, and accommodate the bronchial airways when they branch from the trachea. The bronchial airways terminate in
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * M ...
which make up the functional tissue ( parenchyma) of the lung, and veins, arteries, nerves, and
lymphatic vessel The lymphatic vessels (or lymph vessels or lymphatics) are thin-walled vessels (tubes), structured like blood vessels, that carry lymph. As part of the lymphatic system, lymph vessels are complementary to the cardiovascular system. Lymph ves ...
s. The trachea and bronchi have plexuses of
lymph capillaries Lymph capillaries or lymphatic capillaries are tiny, thin-walled microvessels located in the spaces between cells (except in the central nervous system and non-vascular tissues) which serve to drain and process extracellular fluid. Upon enterin ...
in their mucosa and submucosa. The smaller bronchi have a single layer of lymph capillaries, and they are absent in the alveoli. The lungs are supplied with the largest lymphatic drainage system of any other organ in the body. Each lung is surrounded by a serous membrane of visceral pleura, which has an underlying layer of loose connective tissue attached to the substance of the lung.


Connective tissue

The connective tissue of the lungs is made up of
elastic Elastic is a word often used to describe or identify certain types of elastomer, elastic used in garments or stretchable fabrics. Elastic may also refer to: Alternative name * Rubber band, ring-shaped band of rubber used to hold objects togethe ...
and
collagen fibres Type I collagen is the most abundant collagen of the human body. It forms large, eosinophilic fibers known as collagen fibers. It is present in scar tissue, the end product when tissue heals by repair, as well as tendons, ligaments, the endomys ...
that are interspersed between the capillaries and the alveolar walls.
Elastin Elastin is a protein that in humans is encoded by the ''ELN'' gene. Elastin is a key component of the extracellular matrix in gnathostomes (jawed vertebrates). It is highly elastic and present in connective tissue allowing many tissues in the bo ...
is the key
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
of the
extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide s ...
and is the main component of the
elastic fibres Elastic fibers (or yellow fibers) are an essential component of the extracellular matrix composed of bundles of proteins (elastin) which are produced by a number of different cell types including fibroblasts, endothelial, smooth muscle, and ai ...
. Elastin gives the necessary elasticity and resilience required for the persistent stretching involved in breathing, known as
lung compliance Lung compliance, or pulmonary compliance, is a measure of the lung's ability to stretch and expand (distensibility of elastic tissue). In clinical practice it is separated into two different measurements, static compliance and dynamic compliance. ...
. It is also responsible for the elastic recoil needed. Elastin is more concentrated in areas of high stress such as the openings of the alveoli, and alveolar junctions. The connective tissue links all the alveoli to form the lung parenchyma which has a sponge-like appearance. The alveoli have interconnecting air passages in their walls known as the pores of Kohn.


Respiratory epithelium

All of the lower respiratory tract including the trachea, bronchi, and bronchioles is lined with respiratory epithelium. This is a
ciliated The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike proj ...
epithelium interspersed with goblet cells which produce mucin the main component of
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
, ciliated cells, basal cells, and in the terminal bronchioles– club cells with actions similar to basal cells, and
macrophage Macrophages (abbreviated as M φ, MΦ or MP) ( el, large eaters, from Greek ''μακρός'' (') = large, ''φαγεῖν'' (') = to eat) are a type of white blood cell of the immune system that engulfs and digests pathogens, such as cancer ce ...
s. The epithelial cells, and the
submucosal gland Submucosal glands can refer to various racemose exocrine glands of the mucous type. These glands secrete mucus to facilitate the movement of particles along the body's various tubes, such as the throat and intestines. The mucosa is the lining of the ...
s throughout the respiratory tract secrete airway surface liquid (ASL), the composition of which is tightly regulated and determines how well mucociliary clearance works. Pulmonary neuroendocrine cells are found throughout the respiratory epithelium including the alveolar epithelium, though they only account for around 0.5 per cent of the total epithelial population. PNECs are innervated airway epithelial cells that are particularly focused at airway junction points. These cells can produce serotonin, dopamine, and norepinephrine, as well as polypeptide products. Cytoplasmic processes from the pulmonary neuroendocrine cells extend into the airway lumen where they may sense the composition of inspired gas.


Bronchial airways

In the bronchi there are incomplete tracheal rings of
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck ...
and smaller plates of cartilage that keep them open. Bronchioles are too narrow to support cartilage and their walls are of
smooth muscle Smooth muscle is an involuntary non- striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit ...
, and this is largely absent in the narrower respiratory bronchioles which are mainly just of epithelium. The absence of cartilage in the terminal bronchioles gives them an alternative name of ''membranous bronchioles''.


Respiratory zone

The conducting zone of the respiratory tract ends at the terminal bronchioles when they branch into the respiratory bronchioles. This marks the beginning of the terminal respiratory unit called the acinus which includes the respiratory bronchioles, the alveolar ducts, alveolar sacs, and alveoli. An acinus measures up to 10 mm in diameter. A primary pulmonary lobule is that part of the acinus that includes the alveolar ducts, sacs, and alveoli but does not include the respiratory bronchioles. The unit described as the secondary pulmonary lobule is the lobule most referred to as the pulmonary lobule or respiratory lobule. This lobule is a discrete unit that is the smallest component of the lung that can be seen without aid. The secondary pulmonary lobule is likely to be made up of between 30 and 50 primary lobules. The lobule is supplied by a terminal bronchiole that branches into respiratory bronchioles. The respiratory bronchioles supply the alveoli in each acinus and is accompanied by a pulmonary artery branch. Each lobule is enclosed by an interlobular septa. Each acinus is incompletely separated by an interlobular septa. The respiratory bronchiole gives rise to the alveolar ducts that lead to the alveolar sacs, which contain two or more alveoli. The walls of the alveoli are extremely thin allowing a fast rate of
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical ...
. The alveoli have interconnecting small air passages in their walls known as the pores of Kohn.


Alveoli

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 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. Type II are larger and they line the alveoli and produce and secrete epithelial lining fluid, and lung surfactant. Type II cells are able to divide and differentiate to Type I cells. The alveolar macrophages have an important role in the
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
. They remove substances which deposit in the alveoli including loose red blood cells that have been forced out from blood vessels.


Microbiota

There is a large presence of
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s in the lungs known as the lung microbiota that interacts with the airway epithelial cells; an interaction of probable importance in maintaining homeostasis. The microbiota is complex and dynamic in healthy people, and altered in diseases such as
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, co ...
and COPD. For example significant changes can take place in COPD following infection with rhinovirus. Fungal genera that are commonly found as mycobiota in the microbiota include '' Candida'', '' Malassezia'', ''
Saccharomyces ''Saccharomyces'' is a genus of fungi that includes many species of yeasts. ''Saccharomyces'' is from Greek σάκχαρον (sugar) and μύκης (fungus) and means ''sugar fungus''. Many members of this genus are considered very important in f ...
'', and '' Aspergillus''.


Respiratory tract

The lower respiratory tract is part of the
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies g ...
, and consists of the
trachea The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air- breathing animals with lungs. The trachea extends from t ...
and the structures below this including the lungs. The trachea receives air from the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its st ...
and travels down to a place where it splits (the carina) into a right and left primary 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 Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * M ...
, 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 from the blood into the lungs to be breathed out. Estimates of the total surface area of lungs vary from ; although this is often quoted in textbooks and the media being "the size of a tennis court", it is actually less than half the size of a singles court. The bronchi in the conducting zone are reinforced with
hyaline cartilage Hyaline cartilage is the glass-like (hyaline) and translucent cartilage found on many joint surfaces. It is also most commonly found in the ribs, nose, larynx, and trachea. Hyaline cartilage is pearl-gray in color, with a firm consistency and h ...
in order to hold open the airways. The bronchioles have no cartilage and are surrounded instead by
smooth muscle Smooth muscle is an involuntary non- striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit ...
. Air is warmed to , humidified and cleansed by the conducting zone. Particles from the air being removed by the
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike proje ...
on the respiratory epithelium lining the passageways, in a process called mucociliary clearance. Pulmonary stretch receptors in the smooth muscle of the airways initiate a reflex known as the
Hering–Breuer reflex The Hering–Breuer inflation reflex, named for Josef Breuer and Ewald Hering, is a reflex triggered to prevent the over-inflation of the lung. Pulmonary stretch receptors present on the wall of bronchi and bronchioles of the airways respond to exce ...
that prevents the lungs from over-inflation, during forceful inspiration.


Blood supply

The lungs have a dual blood supply provided by a
bronchial A bronchus is a passage or airway in the lower respiratory tract that conducts air into the lungs. The first or primary bronchi pronounced (BRAN-KAI) to branch from the trachea at the carina are the right main bronchus and the left main bronchus ...
and a pulmonary circulation. The bronchial circulation supplies oxygenated blood to the airways of the lungs, through the
bronchial arteries In human anatomy, the bronchial arteries supply the lungs with nutrition and oxygenated blood. Although there is much variation, there are usually two bronchial arteries that run to the left lung, and one to the right lung and are a vital part ...
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% of the total blood volume of the entire circulatory system. This quantity can easily fluctuate from between one-half and twice the normal volume. Also, in the event of blood loss through hemorrhage, blood from the lungs can partially compensate by automatically transferring to the systemic circulation.


Nerve supply

The lungs are supplied by nerves of the
autonomic nervous system The autonomic nervous system (ANS), formerly referred to as the vegetative nervous system, is a division of the peripheral nervous system that supplies internal organs, smooth muscle and glands. The autonomic nervous system is a control system t ...
. Input from the
parasympathetic nervous system The parasympathetic nervous system (PSNS) is one of the three divisions of the autonomic nervous system, the others being the sympathetic nervous system and the enteric nervous system. The enteric nervous system is sometimes considered part o ...
occurs via the
vagus nerve The vagus nerve, also known as the tenth cranial nerve, cranial nerve X, or simply CN X, is a cranial nerve that interfaces with the parasympathetic control of the heart, lungs, and digestive tract. It comprises two nerves—the left and righ ...
. When stimulated by
acetylcholine Acetylcholine (ACh) is an organic chemical that functions in the brain and body of many types of animals (including humans) as a neurotransmitter. Its name is derived from its chemical structure: it is an ester of acetic acid and choline. Par ...
, 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 Norepinephrine (NE), also called noradrenaline (NA) or noradrenalin, is an organic chemical in the catecholamine family that functions in the brain and body as both a hormone and neurotransmitter. The name "noradrenaline" (from Latin '' ad ...
acting on the beta 2 adrenoceptors in the respiratory tract, which causes
bronchodilation A bronchodilator or broncholytic (although the latter occasionally includes secretory inhibition as well) is a substance that dilates the bronchi and bronchioles, decreasing resistance in the respiratory airway and increasing airflow to the lung ...
. The action of breathing takes place because of nerve signals sent by the
respiratory center The respiratory center is located in the medulla oblongata and pons, in the brainstem. The respiratory center is made up of three major respiratory groups of neurons, two in the medulla and one in the pons. In the medulla they are the dorsal ...
in the brainstem, along the
phrenic nerve The phrenic nerve is a mixed motor/sensory nerve which originates from the C3-C5 spinal nerves in the neck. The nerve is important for breathing because it provides exclusive motor control of the diaphragm, the primary muscle of respiration. ...
from the cervical plexus to the diaphragm.


Variation

The lobes of the lung are subject to
anatomical variation An anatomical variation, anatomical variant, or anatomical variability is a presentation of body structure with morphological features different from those that are typically described in the majority of individuals. Anatomical variations are categ ...
s. A horizontal interlobar fissure was found to be incomplete in 25% of right lungs, or even absent in 11% of all cases. An accessory fissure was also found in 14% and 22% of left and right lungs, respectively. An oblique fissure was found to be incomplete in 21% to 47% of left lungs. In some cases a fissure is absent, or extra, resulting in a right lung with only two lobes, or a left lung with three lobes. A variation in the airway branching structure has been found specifically in the central airway branching. This variation is associated with the development of COPD in adulthood.


Development

The development of the human lungs arise from the
laryngotracheal groove The laryngotracheal groove is a precursor for the larynx and trachea. The rudiment of the respiratory organs appears as a median longitudinal groove in the ventral wall of the pharynx. The groove deepens and its lips fuse to form a septum which gr ...
and develop to maturity over several weeks in the foetus and for several years following birth. The larynx,
trachea The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air- breathing animals with lungs. The trachea extends from t ...
, bronchi and lungs that make up the respiratory tract, begin to form during the fourth week of
embryogenesis An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
from the
lung bud The lung bud sometimes referred to as the respiratory bud forms from the respiratory diverticulum, an embryological endodermal structure that develops into the respiratory tract organs such as the larynx, trachea, bronchi and lungs. It arises from ...
which appears ventrally to the caudal portion of the foregut. The respiratory tract has a branching structure, and is also known as the respiratory 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 proteinsonic 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 Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * M ...
. 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 Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a ...
will take place, together with the type II alveolar cells that secrete
pulmonary surfactant Pulmonary surfactant is a surface-active complex of phospholipids and proteins formed by type II alveolar cells. The proteins and lipids that make up the surfactant have both hydrophilic and hydrophobic regions. By adsorbing to the air-water in ...
, appear. The surfactant reduces the
surface tension Surface tension is the tendency of liquid surfaces at rest to shrink into the minimum surface area possible. Surface tension is what allows objects with a higher density than water such as razor blades and insects (e.g. water striders) t ...
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.


Vitamin A deficiency

The developing lung is particularly vulnerable to changes in the levels of vitamin A. Vitamin A deficiency has been linked to changes in the epithelial lining of the lung and in the lung parenchyma. This can disrupt the normal physiology of the lung and predispose to respiratory diseases. Severe nutritional deficiency in vitamin A results in a reduction in the formation of the alveolar walls (septa) and to notable changes in the respiratory epithelium; alterations are noted in the extracellular matrix and in the protein content of the basement membrane. The extracellular matrix maintains lung elasticity; the basement membrane is associated with alveolar epithelium and is important in the blood-air barrier. The deficiency is associated with functional defects and disease states. Vitamin A is crucial in the development of the alveoli which continues for several years after birth.


After birth

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 The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
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 begin 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.


Function


Gas exchange

The major function of the lungs is
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a ...
between the lungs and the blood. The alveolar and pulmonary capillary gases equilibrate across the thin blood–air barrier. This thin membrane (about 0.5 –2 μm thick) is folded into about 300 million alveoli, providing an extremely large surface area (estimates varying between 70 and 145 m2) for gas exchange to occur. The lungs are not capable of expanding to breathe on their own, 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 Diaphragm may refer to: Anatomy * Thoracic diaphragm, a thin sheet of muscle between the thorax and the abdomen * Pelvic diaphragm or pelvic floor, a pelvic structure * Urogenital diaphragm or triangular ligament, a pelvic structure Other * Diap ...
, and the intercostal muscles which pull the rib cage upwards as shown in the diagram. During
breathing out Breathing (or ventilation) is the process of moving air into and from the lungs to facilitate gas exchange with the internal environment, mostly to flush out carbon dioxide and bring in oxygen. All aerobic creatures need oxygen for cellular ...
the muscles relax, returning the lungs to their resting position. 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 in exertion, a large number of accessory muscles in the neck and abdomen are recruited, that during exhalation pull the ribcage down, decreasing the volume of the thoracic cavity. The FRC is now decreased, but since the lungs cannot be emptied completely there is still about a litre of residual air left. Lung function testing is carried out to evaluate lung volumes and capacities.


Protection

The lungs possess several characteristics which protect against infection. The respiratory tract is lined by respiratory epithelium or respiratory mucosa, with hair-like projections called
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike proje ...
that beat rhythmically and carry
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
. 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 the airways, 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 Goblet cells are simple columnar epithelial cells that secrete gel-forming mucins, like mucin 5AC. The goblet cells mainly use the merocrine method of secretion, secreting vesicles into a duct, but may use apocrine methods, budding off their s ...
secrete mucus which also contains several antimicrobial compounds such as defensins, antiproteases, and
antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...
s. A rare type of specialised cell called a pulmonary ionocyte that is suggested may regulate mucus viscosity has been described. In addition, the lining of the lung also contains
macrophage Macrophages (abbreviated as M φ, MΦ or MP) ( el, large eaters, from Greek ''μακρός'' (') = large, ''φαγεῖν'' (') = to eat) are a type of white blood cell of the immune system that engulfs and digests pathogens, such as cancer ce ...
s, immune cells which engulf and destroy debris and microbes that enter the lung in a process known as phagocytosis; and
dendritic cell Dendritic cells (DCs) are antigen-presenting cells (also known as ''accessory cells'') of the mammalian immune system. Their main function is to process antigen material and present it on the cell surface to the T cells of the immune system. Th ...
s which present antigens to activate components of the
adaptive immune system The adaptive immune system, also known as the acquired immune system, is a subsystem of the immune system that is composed of specialized, systemic cells and processes that eliminate pathogens or prevent their growth. The acquired immune system ...
such as
T cell A T cell is a type of lymphocyte. T cells are one of the important white blood cells of the immune system and play a central role in the adaptive immune response. T cells can be distinguished from other lymphocytes by the presence of a T-cell r ...
s 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 In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...
and behind the mouth in the oropharynx, and larger particles are trapped in nasal hair after inhalation.


Other

In addition to their function in respiration, the lungs have a number of other functions. They are involved in maintaining
homeostasis In biology, homeostasis ( British also homoeostasis) (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physical, and chemical conditions maintained by living systems. This is the condition of optimal functioning for the organism and ...
, helping in the regulation of
blood pressure Blood pressure (BP) is the pressure of circulating blood against the walls of blood vessels. Most of this pressure results from the heart pumping blood through the circulatory system. When used without qualification, the term "blood pressure ...
as part of the
renin–angiotensin system The renin–angiotensin system (RAS), or renin–angiotensin–aldosterone system (RAAS), is a hormone system that regulates blood pressure, fluid and electrolyte balance, and systemic vascular resistance. When renal blood flow is reduced, ...
. The inner lining of the blood vessels secretes
angiotensin-converting enzyme Angiotensin-converting enzyme (), or ACE, is a central component of the renin–angiotensin system (RAS), which controls blood pressure by regulating the volume of fluids in the body. It converts the hormone angiotensin I to the active vasoconst ...
(ACE) an
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
that catalyses the conversion of
angiotensin I Angiotensin is a peptide hormone that causes vasoconstriction and an increase in blood pressure. It is part of the renin–angiotensin system, which regulates blood pressure. Angiotensin also stimulates the release of aldosterone from the adre ...
to
angiotensin II Angiotensin is a peptide hormone that causes vasoconstriction and an increase in blood pressure. It is part of the renin–angiotensin system, which regulates blood pressure. Angiotensin also stimulates the release of aldosterone from the adr ...
. The lungs are involved in the blood's acid–base homeostasis by expelling
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
when breathing. The lungs also serve a protective role. Several blood-borne substances, such as a few types of prostaglandins, leukotrienes,
serotonin Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and va ...
and
bradykinin Bradykinin (BK) (Greek brady-, slow; -kinin, kīn(eîn) to move) is a peptide that promotes inflammation. It causes arterioles to dilate (enlarge) via the release of prostacyclin, nitric oxide, and endothelium-derived hyperpolarizing factor an ...
, 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
vein Veins are blood vessels in humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated ...
s and prevent them from entering arteries and causing
stroke A stroke is a disease, medical condition in which poor cerebral circulation, blood flow to the brain causes cell death. There are two main types of stroke: brain ischemia, ischemic, due to lack of blood flow, and intracranial hemorrhage, hemorr ...
s. The lungs also play a pivotal role in speech by providing air and airflow for the creation of vocal sounds, and other paralanguage
communications Communication (from la, communicare, meaning "to share" or "to be in relation with") is usually defined as the transmission of information. The term may also refer to the message communicated through such transmissions or the field of inquir ...
such as
sighs Paralanguage, also known as vocalics, is a component of meta-communication that may modify meaning, give nuanced meaning, or convey emotion, by using techniques such as prosody, pitch, volume, intonation, etc. It is sometimes defined as relatin ...
and
gasp Paralanguage, also known as vocalics, is a component of meta-communication that may modify meaning, give nuanced meaning, or convey emotion, by using techniques such as prosody, pitch, volume, intonation, etc. It is sometimes defined as relatin ...
s. Research suggests a role of the lungs in the production of blood platelets.


Gene and protein expression

About 20,000 protein coding genes are expressed in human cells and almost 75% of these genes are expressed in the normal lung. A little less than 200 of these genes are more specifically expressed in the lung with less than 20 genes being highly lung specific. The highest expression of lung specific proteins are different surfactant proteins, such as
SFTPA1 Surfactant protein A1 (SP-A1), also known as Pulmonary surfactant-associated protein A1 (PSP-A) is a protein that in humans is encoded by the ''SFTPA1'' gene. Summary SP-A1 is primarily synthesised in type II alveolar cells in the lung, as pa ...
, SFTPB and
SFTPC Surfactant protein C (SP-C), is one of the pulmonary surfactant proteins. In humans this is encoded by the ''SFTPC'' gene. It is a membrane protein. Structure SFTPC is a 197-residue protein made up of two halves: a unique N-terminal propepti ...
, and napsin, expressed in type II pneumocytes. Other proteins with elevated expression in the lung are the dynein protein DNAH5 in ciliated cells, and the secreted SCGB1A1 protein in mucus-secreting goblet cells of the airway mucosa.


Clinical significance

Lungs can be affected by a number of diseases and disorders. Pulmonology is the
medical speciality A medical specialty is a branch of medical practice that is focused on a defined group of patients, diseases, skills, or philosophy. Examples include those branches of medicine that deal exclusively with children (paediatrics), cancer (oncology), ...
that deals with respiratory diseases involving the lungs and
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies g ...
. Cardiothoracic surgery deals with
surgery Surgery ''cheirourgikē'' (composed of χείρ, "hand", and ἔργον, "work"), via la, chirurgiae, meaning "hand work". is a medical specialty that uses operative manual and instrumental techniques on a person to investigate or treat a pa ...
of the lungs including lung volume reduction surgery, lobectomy,
pneumectomy A pneumonectomy (or pneumectomy) is a surgery, surgical procedure to remove a lung first successfully done in 1933 by Dr. Evarts Graham. This is not to be confused with a lobectomy or segmentectomy, which only removes one part of the lung. Ther ...
and lung transplantation.


Inflammation and infection

Inflammatory conditions of the lung tissue are
pneumonia Pneumonia is an inflammatory condition of the lung primarily affecting the small air sacs known as alveoli. Symptoms typically include some combination of productive or dry cough, chest pain, fever, and difficulty breathing. The severit ...
, of the respiratory tract are bronchitis and bronchiolitis, and of the pleurae surrounding the lungs pleurisy. Inflammation is usually caused by
infection An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable di ...
s due to
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
or
virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsk ...
es. When the lung tissue is inflamed due to other causes it is called pneumonitis. One major cause of bacterial pneumonia is
tuberculosis Tuberculosis (TB) is an infectious disease usually caused by '' Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, ...
. 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. Alcohol affects the lungs and can cause inflammatory alcoholic lung disease. Acute exposure to alcohol stimulates the beating of
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike proje ...
in the respiratory epithelium. However, chronic exposure has the effect of desensitising the ciliary response which reduces mucociliary clearance (MCC). MCC is an innate defense system protecting against pollutants and pathogens, and when this is disrupted the numbers of alveolar macrophages are decreased. A subsequent inflammatory response is the release of
cytokine Cytokines are a broad and loose category of small proteins (~5–25 kDa) important in cell signaling. Cytokines are peptides and cannot cross the lipid bilayer of cells to enter the cytoplasm. Cytokines have been shown to be involved in au ...
s. Another consequence is the susceptibility to infection.


Blood-supply changes

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 D-dimer (or D dimer) is a fibrin degradation product (or FDP), a small protein fragment present in the blood after a blood clot is degraded by fibrinolysis. It is so named because it contains two D fragments of the fibrin protein joined by a cros ...
. 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 Granulomatosis with polyangiitis (GPA), previously known as Wegener's granulomatosis (WG), is a rare long-term systemic disorder that involves the formation of granulomas and inflammation of blood vessels (vasculitis). It is a form of vasculitis ...
, which causes inflammation of the small blood vessels of the lungs and kidneys. A
lung contusion The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side ...
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 A surgical drain is a tube used to remove pus, blood or other fluids from a wound, body cavity, or organ. They are commonly placed by surgeons or interventional radiologists after procedures or some types of injuries, but they can also be used a ...
until the underlying cause is identified and treated.


Obstructive lung diseases

Asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, co ...
,
bronchiectasis Bronchiectasis is a disease in which there is permanent enlargement of parts of the airways of the lung. Symptoms typically include a chronic cough with mucus production. Other symptoms include shortness of breath, coughing up blood, and chest ...
, and
chronic obstructive pulmonary disease Chronic obstructive pulmonary disease (COPD) is a type of progressive lung disease characterized by long-term respiratory symptoms and airflow limitation. The main symptoms include shortness of breath and a cough, which may or may not produce ...
(COPD) that includes chronic bronchitis, and
emphysema Emphysema, or pulmonary emphysema, is a lower respiratory tract disease, characterised by air-filled spaces ( pneumatoses) in the lungs, that can vary in size and may be very large. The spaces are caused by the breakdown of the walls of the alv ...
, are all obstructive lung diseases characterised by
airway obstruction Airway obstruction is a blockage of respiration in the airway that hinders the free flow of air. It can be broadly classified into being either in the upper airway (UPA) or lower airway (LOA). Airway obstruction is a life-threatening condition ...
. 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 Pulmonary function testing (PFT) is a complete evaluation of the respiratory system including patient history, physical examinations, and tests of pulmonary function. The primary purpose of pulmonary function testing is to identify the severity ...
such as spirometry. Many obstructive lung diseases are managed by avoiding triggers (such as
dust mite House dust mites (HDM, or simply dust mites) are various species of acariform mites belonging to the family Pyroglyphidae that are found in association with dust in dwellings. They are known for causing allergies. Biology Species The curren ...
s or
smoking Smoking is a practice in which a substance is burned and the resulting smoke is typically breathed in to be tasted and absorbed into the bloodstream. Most commonly, the substance used is the dried leaves of the tobacco plant, which have b ...
), with symptom control such as bronchodilators, and with suppression of inflammation (such as through
corticosteroid Corticosteroids are a class of steroid hormones that are produced in the adrenal cortex of vertebrates, as well as the synthetic analogues of these hormones. Two main classes of corticosteroids, glucocorticoids and mineralocorticoids, are inv ...
s) in severe cases. A common cause of chronic bronchitis, and emphysema, is smoking; and common causes of
bronchiectasis Bronchiectasis is a disease in which there is permanent enlargement of parts of the airways of the lung. Symptoms typically include a chronic cough with mucus production. Other symptoms include shortness of breath, coughing up blood, and chest ...
include severe infections and cystic fibrosis. The definitive cause of
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, co ...
is not yet known. The breakdown of alveolar tissue, often as a result of tobacco-smoking leads to emphysema, which can become severe enough to develop into COPD. Elastase breaks down the
elastin Elastin is a protein that in humans is encoded by the ''ELN'' gene. Elastin is a key component of the extracellular matrix in gnathostomes (jawed vertebrates). It is highly elastic and present in connective tissue allowing many tissues in the bo ...
in the lung's connective tissue that can also result in emphysema. Elastase is inhibited by the
acute-phase protein Acute-phase proteins (APPs) are a class of proteins whose concentrations in blood plasma either increase (positive acute-phase proteins) or decrease (negative acute-phase proteins) in response to inflammation. This response is called the ''acute- ...
, alpha-1 antitrypsin, and when there is a deficiency in this, emphysema can develop. With persistent stress from smoking, the airway basal cells become disarranged and lose their regenerative ability needed to repair the epithelial barrier. The disorganised basal cells are seen to be responsible for the major airway changes that are characteristic of COPD, and with continued stress can undergo a malignant transformation. Studies have shown that the initial development of emphysema is centred on the early changes in the airway epithelium of the small airways. Basal cells become further deranged in a smoker's transition to clinically defined COPD.


Restrictive lung diseases

Some types of chronic lung diseases are classified as
restrictive lung disease Restrictive lung diseases are a category of extrapulmonary, pleural, or parenchymal respiratory diseases that restrict lung expansion, resulting in a decreased lung volume, an increased work of breathing, and inadequate ventilation and/or oxygena ...
, 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 Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tissue is ...
. This can be due to a large variety of occupational lung diseases such as
Coalworker's pneumoconiosis Coal workers' pneumoconiosis (CWP), also known as black lung disease or black lung, is an occupational type of pneumoconiosis caused by long-term exposure to coal dust. It is common in coal miners and others who work with coal. It is similar to b ...
,
autoimmune disease An autoimmune disease is a condition arising from an abnormal immune response to a functioning body part. At least 80 types of autoimmune diseases have been identified, with some evidence suggesting that there may be more than 100 types. Nearly a ...
s or more rarely to a reaction to
medication A medication (also called medicament, medicine, pharmaceutical drug, medicinal drug or simply drug) is a drug used to diagnose, cure, treat, or prevent disease. Drug therapy ( pharmacotherapy) is an important part of the medical field and ...
. Severe respiratory disorders, where spontaneous breathing is not enough to maintain life, may need the use of
mechanical ventilation Mechanical ventilation, assisted ventilation or intermittent mandatory ventilation (IMV), is the medical term for using a machine called a ventilator to fully or partially provide artificial ventilation. Mechanical ventilation helps move a ...
to ensure an adequate supply of air.


Cancers

Lung cancer Lung cancer, also known as lung carcinoma (since about 98–99% of all lung cancers are carcinomas), is a malignant lung tumor characterized by uncontrolled cell growth in tissues of the lung. Lung carcinomas derive from transformed, mali ...
can either arise directly from lung tissue or as a result of
metastasis Metastasis is a pathogenic agent's spread from an initial or primary site to a different or secondary site within the host's body; the term is typically used when referring to metastasis by a cancerous tumor. The newly pathological sites, the ...
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 Smoking is a practice in which a substance is burned and the resulting smoke is typically breathed in to be tasted and absorbed into the bloodstream. Most commonly, the substance used is the dried leaves of the tobacco plant, which have b ...
. Once a cancer is identified it is
staged ''Staged'' is a British television comedy series, set during the COVID-19 pandemic in the United Kingdom and primarily filmed using video-conferencing technology. The first series premiered on 10 June 2020 on BBC One, and the second series pre ...
using scans such as a CT scan and a sample of tissue from a
biopsy A biopsy is a medical test commonly performed by a surgeon, interventional radiologist, or an interventional cardiologist. The process involves extraction of sample cells or tissues for examination to determine the presence or extent of a dise ...
is taken. Cancers may be treated surgically by removing the tumour, the use of
radiotherapy Radiation therapy or radiotherapy, often abbreviated RT, RTx, or XRT, is a therapy using ionizing radiation, generally provided as part of cancer treatment to control or kill malignant cells and normally delivered by a linear accelerator. Rad ...
,
chemotherapy Chemotherapy (often abbreviated to chemo and sometimes CTX or CTx) is a type of cancer treatment that uses one or more anti-cancer drugs (chemotherapeutic agents or alkylating agents) as part of a standardized chemotherapy regimen. Chemother ...
or a combination, or with the aim of symptom control.
Lung cancer screening Lung cancer screening refers to cancer screening strategies used to identify early lung cancers before they cause symptoms, at a point where they are more likely to be curable. Lung cancer screening is critically important because of the inciden ...
is being recommended in the United States for high-risk populations.


Congenital disorders

Congenital disorder A birth defect, also known as a congenital disorder, is an abnormal condition that is present at birth regardless of its cause. Birth defects may result in disabilities that may be physical, intellectual, or developmental. The disabilities c ...
s include cystic fibrosis,
pulmonary hypoplasia Pulmonary hypoplasia is incomplete development of the lungs, resulting in an abnormally low number or size of bronchopulmonary segments or alveoli. A congenital malformation, it most often occurs secondary to other fetal abnormalities that i ...
(an incomplete development of the lungs) congenital diaphragmatic hernia, and
infant respiratory distress syndrome Infantile respiratory distress syndrome (IRDS), also called respiratory distress syndrome of newborn, or increasingly surfactant deficiency disorder (SDD), and previously called hyaline membrane disease (HMD), is a syndrome in premature infants ...
caused by a deficiency in lung surfactant. An azygos lobe is a congenital
anatomical variation An anatomical variation, anatomical variant, or anatomical variability is a presentation of body structure with morphological features different from those that are typically described in the majority of individuals. Anatomical variations are categ ...
which though usually without effect can cause problems in thoracoscopic procedures.


Pleural space pressure

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 thoracic wall or chest wall is the boundary of the thoracic cavity. Structure The bony skeletal part of the thoracic wall is the rib cage, and the rest is made up of muscle, skin, and fasciae. The chest wall has 10 layers, namely (from sup ...
. 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, scuba divers ascending while holding their breath with their lungs fully inflated can cause air sacs (
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * M ...
) to burst and leak high pressure air into the pleural space.


Examination

As part of a
physical examination In a physical examination, medical examination, or clinical examination, a medical practitioner examines a patient for any possible medical signs or symptoms of a medical condition. It generally consists of a series of questions about the pati ...
in response to respiratory symptoms of
shortness of breath Shortness of breath (SOB), also medically known as dyspnea (in AmE) or dyspnoea (in BrE), is an uncomfortable feeling of not being able to breathe well enough. The American Thoracic Society defines it as "a subjective experience of breathing di ...
, and cough, a lung examination may be carried out. This exam includes palpation and
auscultation Auscultation (based on the Latin verb ''auscultare'' "to listen") is listening to the internal sounds of the body, usually using a stethoscope. Auscultation is performed for the purposes of examining the circulatory and respiratory systems (he ...
. The areas of the lungs that can be listened to using a stethoscope are called the
lung fields A respiratory examination, or lung examination, is performed as part of a physical examination, in response to respiratory symptoms such as shortness of breath, cough, or chest pain, and is often carried out with a cardiac examination. The four ...
, and these are the posterior, lateral, and anterior lung fields. The posterior fields can be listened to from the back and include: the lower lobes (taking up three quarters of the posterior fields); the anterior fields taking up the other quarter; and the lateral fields under the axillae, the left axilla for the lingual, the right axilla for the middle right lobe. The anterior fields can also be auscultated from the front. An area known as the
triangle of auscultation The triangle of auscultation is a relative thinning of the musculature of the back, situated along the medial border of the scapula which allows for improved listening to the lungs. Boundaries It has the following boundaries: * medially, by th ...
is an area of thinner musculature on the back which allows improved listening. Abnormal breathing sounds heard during a lung exam can indicate the presence of a lung condition;
wheezing A wheeze is a continuous, coarse, whistling sound produced in the respiratory airways during breathing. For wheezes to occur, some part of the respiratory tree must be narrowed or obstructed (for example narrowing of the lower respiratory tract ...
for example is commonly associated with
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, co ...
and COPD.


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. Lung capacity can be increased by physical training as much as 40% but the effect may be modified by exposure to air pollution. 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 Spirometry (meaning ''the measuring of breath'') is the most common of the pulmonary function tests (PFTs). It measures lung function, specifically the amount (volume) and/or speed (flow) of air that can be inhaled and exhaled. Spirometry is he ...
(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.


Other animals


Birds

The lungs of birds are relatively small, but are connected to 8 or 9
air sacs Air sacs are spaces within an organism where there is the constant presence of air. Among modern animals, birds possess the most air sacs (9–11), with their extinct dinosaurian relatives showing a great increase in the pneumatization (presence o ...
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 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; these, like the alveoli in other lungs, are the site of
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a ...
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''.


Reptiles

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,
snake Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more ...
s, 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 lungs similar to those of birds, providing a 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 ''Tupuxuara'' is a genus of large, crested, and toothless pterodactyloid pterosaur from the Early Cretaceous period (Albian stage) of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million years ago. ''Tupuxu ...
, and azhdarchoids, the hindlimbs. Reptilian lungs typically receive air via expansion and contraction of the ribs driven by axial muscles and buccal pumping.
Crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest livin ...
s 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.
Turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s, which are unable to move their ribs, instead use their forelimbs and pectoral girdle to force air in and out of the lungs.


Amphibians

The lungs of most
frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s 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 The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, forcing the air into the lungs. Due to the possibility of respiration across the skin combined with small size, all known lungless
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s are amphibians. The majority of salamander species are lungless salamanders, which respirate through their skin and tissues lining their mouth. This necessarily restricts 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 ''Atretochoana eiselti'' is a species of caecilian originally known only from two preserved specimens discovered by Sir Graham Hales in the Brazilian rainforest, while on an expedition with Sir Brian Doll in the late 1800s, but rediscovered in 2 ...
'', a caecilian. The lungs of amphibians typically have a few narrow internal walls (
septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five c ...
) of soft tissue around the outer walls, increasing the respiratory surface area and giving the lung a honeycomb 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.


Lungfish

The lungs of
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
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 esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to t ...
. 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.


Invertebrates

Some
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s have lung-like structures that serve a similar respiratory purpose as, but are not evolutionarily related to, vertebrate lungs. Some arachnids, such as
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s and
scorpion Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always en ...
s, have structures called
book lung A book lung is a type of respiration organ used for atmospheric gas exchange that is present in many arachnids, such as scorpions and spiders. Each of these organs is located inside an open ventral abdominal, air-filled cavity (atrium) and conn ...
s 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 The coconut crab (''Birgus latro'') is a species of terrestrial hermit crab, also known as the robber crab or palm thief. It is the largest terrestrial arthropod in the world, with a weight of up to . It can grow to up to in width from the tip ...
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.


Evolutionary origins

The lungs of today's terrestrial
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s and the gas bladders of today's
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
are believed to have evolved from simple sacs, as outpocketings of the
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to t ...
, that allowed early fish to gulp air under oxygen-poor conditions. These outpocketings first arose in the
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...
. In most of the
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
the sacs evolved into closed off gas bladders, while a number of
carp Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
, trout, herring,
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...
, and eels have retained the physostome condition with the sac 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
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s. Thus, the lungs of vertebrates are homologous to the gas bladders of fish (but not to their
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they ar ...
s).


See also

*
Atelectasis Atelectasis is the collapse or closure of a lung resulting in reduced or absent gas exchange. It is usually unilateral, affecting part or all of one lung. It is a condition where the alveoli are deflated down to little or no volume, as distinct ...
* Drowning * Interstitial lung disease *
Liquid breathing Liquid breathing is a form of respiration in which a normally air-breathing organism breathes an oxygen-rich liquid (such as a perfluorocarbon), rather than breathing air. By selecting a liquid that is capable of holding large amounts of ...
*
Lung abscess Lung abscess is a type of liquefactive necrosis of the lung tissue and formation of cavities (more than 2 cm) containing necrotic debris or fluid caused by microbial infection. This pus-filled cavity is often caused by aspiration, which may ...
*
Lung-on-a-chip The lung-on-a-chip is a complex, three-dimensional model of a living, breathing human lung on a microchip. The device is made using human lung and blood vessel cells and it can predict absorption of airborne nanoparticles and mimic the inflammato ...
*
Secarecytosis Secarecytosis is a process involved in the development of a bird's lung cells, before the bird hatches from its egg. It is the processes of cell cutting during attenuation of the tubular epithelium of the developing avian lung. The word ''secarecyto ...
*
List of terms of lung size and activity Following are terms that specify a type of lung size and/or activity. More specific definitions may be found in individual articles. *Eupnea – normal breathing *Apnea – absence of breathing *Bradypnea – decreased breathing rate *Dyspnea or '' ...


References


Further reading


Dr D.R. Johnson: Introductory anatomy, respiratory system
leeds.ac.uk

sln.fi.edu

people.eku.edu


External links


Lung at the Human Protein Atlas
{{DEFAULTSORT:Human Lung Human anatomy by organ Articles containing video clips