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The lungs are the primary
organs In biology, an organ is a collection of tissues joined in a structural unit to serve a common function. In the hierarchy of life, an organ lies between tissue and an organ system. Tissues are formed from same type cells to act together in a f ...
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 ...
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 Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
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 G ...
s and a small number of
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
. In mammals 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 c ...
s, two lungs are located near the
backbone The backbone is the vertebral column of a vertebrate. Arts, entertainment, and media Film * ''Backbone'' (1923 film), a 1923 lost silent film starring Alfred Lunt * ''Backbone'' (1975 film), a 1975 Yugoslavian drama directed by Vlatko Gilić ...
on either side of the
heart The heart is a muscular 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 carbon dioxide to t ...
. 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 as ...
from the
air The atmosphere of Earth is the layer of gases, known collectively as air, retained by Earth's gravity that surrounds the planet and forms its planetary atmosphere. The atmosphere of Earth protects life on Earth by creating pressure allowing f ...
and transfer it into the
bloodstream The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
, 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 trans ...
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 Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
is driven by different muscular systems in different species. Mammals, reptiles 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 The pharyngeal muscles are a group of muscles that form the pharynx, which is posterior to the oral cavity, determining the shape of its lumen, and affecting its sound properties as the primary resonating cavity. The pharyngeal muscles (involunta ...
via
buccal pumping Buccal pumping is "breathing with one's cheeks": a method of ventilation used in respiration in which the animal moves the floor of its mouth in a rhythmic manner that is externally apparent.Brainerd, E. L. (1999). New perspectives on the evolutio ...
, a mechanism still seen in amphibians. In humans, the main muscle of respiration that drives breathing is the diaphragm. The lungs also provide airflow that makes vocal sounds including human speech possible. Humans have two lungs, one on the left and one on the right. They are situated within the
thoracic cavity The thoracic cavity (or chest cavity) is the chamber of the body of vertebrates that is protected by the thoracic wall (rib cage and associated skin, muscle, and fascia). The central compartment of the thoracic cavity is the mediastinum. There ...
of the
chest The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
. 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 The respiratory tract is the subdivision of the respiratory system involved with the process of Respiration (physiology), respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed ...
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 the ...
and branches into the
bronchi 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
bronchiole The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start ...
s, and which receive
air The atmosphere of Earth is the layer of gases, known collectively as air, retained by Earth's gravity that surrounds the planet and forms its planetary atmosphere. The atmosphere of Earth protects life on Earth by creating pressure allowing f ...
breathed in via the
conducting zone The respiratory tract is the subdivision of the respiratory system involved with the process of respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed in through the nose to ...
. The conducting zone ends at the
terminal bronchioles The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start o ...
. These divide into the
respiratory bronchioles The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start o ...
of the
respiratory zone The respiratory tract is the subdivision of the respiratory system involved with the process of respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed in through the nose to t ...
which divide into alveolar ducts that give rise to the
alveolar sac A pulmonary alveolus (plural: alveoli, from Latin ''alveolus'', "little cavity"), also known as an air sac or air space, is one of millions of hollow, distensible cup-shaped cavities in the lungs where oxygen is exchanged for carbon dioxide. Al ...
s that contain the alveoli, 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 called
pleurae The pulmonary pleurae (''sing.'' pleura) are the two opposing layers of serous membrane overlying the lungs and the inside of the surrounding chest walls. The inner pleura, called the visceral pleura, covers the surface of each lung and dips be ...
; the membranes are separated by a film of
pleural fluid The pleural cavity, pleural space, or interpleural space is the potential space between the pleurae of the pleural sac that surrounds each lung. A small amount of serous pleural fluid is maintained in the pleural cavity to enable lubrication be ...
, 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 The pulmonary circulation is a division of the circulatory system in all vertebrates. The circuit begins with deoxygenated blood returned from the body to the right atrium of the heart where it is pumped out from the right ventricle to the lungs ...
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 bronchial circulation is the part of the circulatory system that supplies nutrients and oxygen to the cells that constitute the lungs, as well as carrying waste products away from them. It is complementary to the pulmonary circulation that brin ...
. The tissue of the lungs can be affected by a number of
respiratory disease Respiratory diseases, or lung diseases, are pathological conditions affecting the organs and tissues that make gas exchange difficult in air-breathing animals. They include conditions of the respiratory tract including the trachea, bronchi, bro ...
s, 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 severi ...
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, malign ...
.
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 Bronchitis is inflammation of the bronchi (large and medium-sized airways) in the lungs that causes coughing. Bronchitis usually begins as an infection in the nose, ears, throat, or sinuses. The infection then makes its way down to the bronchi. ...
and emphysema, and can be related to smoking or exposure to harmful substances. A number of
occupational lung disease Occupational lung diseases are work-related, lung conditions that have been caused or made worse by the materials a person is exposed to within the workplace. It includes a broad group of diseases, including occupational asthma, industrial bronch ...
s can be caused by substances such as
coal dust Coal dust is a fine powdered form of which is created by the crushing, grinding, or pulverizing of coal. Because of the brittle nature of coal, coal dust can be created during mining, transportation, or by mechanically handling coal. It is a form ...
, 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 Bronchitis is inflammation of the bronchi (large and medium-sized airways) in the lungs that causes coughing. Bronchitis usually begins as an infection in the nose, ears, throat, or sinuses. The infection then makes its way down to the bronchi. ...
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 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 the power of the ...
''pulmonarius'' (of the lungs) as in pulmonology, or with ''pneumo-'' (from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
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 severi ...
. 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 s ...
and so they do not function to breathe. Blood is also diverted from the lungs through the
ductus arteriosus The ''ductus arteriosus'', also called the ''ductus Botalli'', named after the Italian physiologist Leonardo Botallo, is a blood vessel in the developing fetus connecting the trunk of the pulmonary artery to the proximal descending aorta. It a ...
. 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 The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
on either side of the
heart The heart is a muscular 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 carbon dioxide to t ...
in the
rib cage 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 sem ...
. They are conical in shape with a narrow rounded apex at the top, and a broad concave base that rests on the convex surface of the diaphragm. The apex of the lung extends into the root of the neck, reaching shortly above the level of the sternal end of the
first rib The 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 semi ...
. The lungs stretch from close to the
backbone The backbone is the vertebral column of a vertebrate. Arts, entertainment, and media Film * ''Backbone'' (1923 film), a 1923 lost silent film starring Alfred Lunt * ''Backbone'' (1975 film), a 1975 Yugoslavian drama directed by Vlatko Gilić ...
in the rib cage to the front of the
chest The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
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 Great vessels are the large vessels that bring blood to and from the heart. These are: *Superior vena cava *Inferior vena cava *Pulmonary arteries * Pulmonary veins *Aorta Transposition of the great vessels is a group of congenital heart defec ...
, and the
carina Carina may refer to: Places Australia * Carina, Queensland, a suburb in Brisbane * Carina Heights, Queensland, a suburb in Brisbane * Carina, Victoria, a locality in Mildura Serbia * Carina, Osečina, a village in the Kolubara District ...
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 away ...
s and airways pass into the lungs making up the
root of the lung The root of the lung is a group of structures that emerge at the hilum of each lung, just above the middle of the mediastinal surface and behind the cardiac impression of the lung. It is nearer to the back (posterior border) than the front (ant ...
. There are also bronchopulmonary lymph nodes on the hilum. The lungs are surrounded by the pulmonary pleurae. The pleurae are two
serous membrane The serous membrane (or serosa) is a smooth tissue membrane of mesothelium lining the contents and inner walls of body cavities, which secrete serous fluid to allow lubricated sliding movements between opposing surfaces. The serous membrane ...
s; the outer
parietal pleura The pulmonary pleurae (''sing.'' pleura) are the two opposing layers of serous membrane overlying the lungs and the inside of the surrounding chest walls. The inner pleura, called the visceral pleura, covers the surface of each lung and dips be ...
lines the inner wall of the
rib cage 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 sem ...
and the inner
visceral pleura The pulmonary pleurae (''sing.'' pleura) are the two opposing layers of serous membrane overlying the lungs and the inside of the surrounding chest walls. The inner pleura, called the visceral pleura, covers the surface of each lung and dips b ...
directly lines the surface of the lungs. Between the pleurae is a
potential space In anatomy, a potential space is a space between two adjacent structures that are normally pressed together (directly apposed). Many anatomic spaces are potential spaces, which means that they are potential rather than realized (with their realiz ...
called the
pleural cavity The pleural cavity, pleural space, or interpleural space is the potential space between the pleurae of the pleural sac that surrounds each lung. A small amount of serous pleural fluid is maintained in the pleural cavity to enable lubrication b ...
containing a thin layer of lubricating
pleural fluid The pleural cavity, pleural space, or interpleural space is the potential space between the pleurae of the pleural sac that surrounds each lung. A small amount of serous pleural fluid is maintained in the pleural cavity to enable lubrication be ...
.


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 Variation or Variations may refer to: Science and mathematics * Variation (astronomy), any perturbation of the mean motion or orbit of a planet or satellite, particularly of the moon * Genetic variation, the difference in DNA among individua ...
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 The costal cartilages are bars of hyaline cartilage that serve to prolong the ribs forward and contribute to the elasticity of the walls of the thorax. Costal cartilage is only found at the anterior ends of the ribs, providing medial extension. ...
; on the mediastinal 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 The azygos vein is a vein running up the right side of the thoracic vertebral column draining itself towards the superior vena cava. It connects the systems of superior vena cava and inferior vena cava and can provide an alternative path for blo ...
, and above this is a wide groove for the
superior vena cava The superior vena cava (SVC) is the superior of the two venae cavae, the great venous trunks that return deoxygenated blood from the systemic circulation to the right atrium of the heart. It is a large-diameter (24 mm) short length vein th ...
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 j ...
; behind this, and close to the top of the lung is a groove for the
brachiocephalic artery The brachiocephalic artery (or brachiocephalic trunk or innominate artery) is an artery of the mediastinum that supplies blood to the right arm and the head and neck. It is the first branch of the aortic arch. Soon after it emerges, the brachioce ...
. There is a groove for the esophagus behind the hilum and the
pulmonary ligament The root of the lung is a group of structures that emerge at the hilum of each lung, just above the middle of the mediastinal surface and behind the cardiac impression of the lung. It is nearer to the back (posterior border) than the front (ant ...
, and near the lower part of the esophageal groove is a deeper groove for the
inferior vena cava The inferior vena cava is a large vein that carries the deoxygenated blood from the lower and middle body into the right atrium of the heart. It is formed by the joining of the right and the left common iliac veins, usually at the level of th ...
before it enters the heart.Alt URL
/ref> The weight of the right lung varies between individuals, with a standard
reference range In medicine and health-related fields, a reference range or reference interval is the range or the interval of values that is deemed normal for a physiological measurement in healthy persons (for example, the amount of creatinine in the blood, o ...
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 Costal may refer to: * an adjective related to the rib () in anatomy ** Costal cartilage, a type of cartilage forming bars which serve to prolong the ribs forward ** Costal margin, the medial margin formed by the false ribs ** Costal surface (disa ...
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 In human anatomy, the descending aorta is part of the aorta, the largest artery in the body. The descending aorta begins at the aortic arch and runs down through the chest and abdomen. The descending aorta anatomically consists of two portions o ...
. 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 j ...
. 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 ...
may sit in a wider shallow impression at the base of the lung. The weight of the left lung, by standard
reference range In medicine and health-related fields, a reference range or reference interval is the range or the interval of values that is deemed normal for a physiological measurement in healthy persons (for example, the amount of creatinine in the blood, o ...
, 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 in ...
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 The respiratory tract is the subdivision of the respiratory system involved with the process of Respiration (physiology), respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed ...
, and accommodate the bronchial airways when they branch from the trachea. The bronchial airways terminate in alveoli 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 ve ...
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 enteri ...
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 The serous membrane (or serosa) is a smooth tissue membrane of mesothelium lining the contents and inner walls of body cavities, which secrete serous fluid to allow lubricated sliding movements between opposing surfaces. The serous membrane ...
of
visceral pleura The pulmonary pleurae (''sing.'' pleura) are the two opposing layers of serous membrane overlying the lungs and the inside of the surrounding chest walls. The inner pleura, called the visceral pleura, covers the surface of each lung and dips b ...
, which has an underlying layer of
loose connective tissue Loose connective tissue, sometimes called areolar tissue, is a cellular connective tissue with thin and relatively sparse collagen fibers. Its ground substance occupies more volume than the fibers do. It has a viscous to gel-like consisten ...
attached to the substance of the lung.


Connective tissue

The connective tissue of the lungs is made up of elastic and collagen fibres that are interspersed between the capillaries and the alveolar walls. Elastin 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. Elastin gives the necessary elasticity and resilience required for the persistent stretching involved in breathing, known as lung 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 Respiratory epithelium, or airway epithelium, is a type of ciliated columnar epithelium found lining most of the respiratory tract as respiratory mucosa, where it serves to moisten and protect the airways. It is not present in the vocal cords o ...
. 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 project ...
epithelium interspersed with goblet cells which produce
mucin Mucins () are a family of high molecular weight, heavily glycosylated proteins (glycoconjugates) produced by epithelial tissues in most animals. Mucins' key characteristic is their ability to form gels; therefore they are a key component in most ...
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 bronchiole The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start o ...
s– club cells with actions similar to basal cells, and macrophages. The epithelial cells, and the submucosal glands throughout the respiratory tract secrete airway surface liquid (ASL), the composition of which is tightly regulated and determines how well
mucociliary clearance Mucociliary clearance (MCC), mucociliary transport, or the mucociliary escalator, describes the self-clearing mechanism of the airways in the respiratory system. It is one of the two protective processes for the lungs in removing inhaled particl ...
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 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 the l ...
of cartilage and smaller plates of cartilage that keep them open. Bronchioles are too narrow to support cartilage and their walls are of smooth muscle, and this is largely absent in the narrower
respiratory bronchiole The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start o ...
s 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 sac A pulmonary alveolus (plural: alveoli, from Latin ''alveolus'', "little cavity"), also known as an air sac or air space, is one of millions of hollow, distensible cup-shaped cavities in the lungs where oxygen is exchanged for carbon dioxide. Al ...
s, 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 A pulmonary artery is an artery in the pulmonary circulation that carries deoxygenated blood from the right side of the heart to the lungs. The largest pulmonary artery is the ''main pulmonary artery'' or ''pulmonary trunk'' from the heart, and ...
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 chemica ...
. 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 A pulmonary alveolus (plural: alveoli, from Latin ''alveolus'', "little cavity"), also known as an air sac or air space, is one of millions of hollow, distensible cup-shaped cavities in the lungs where oxygen is exchanged for carbon dioxide. Al ...
and an
alveolar macrophage An alveolar macrophage, pulmonary macrophage, (or dust cell) is a type of macrophage, a professional phagocyte, found in the airways and at the level of the alveoli in the lungs, but separated from their walls. Activity of the alveolar macropha ...
. 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 Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
"), 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 macrophage An alveolar macrophage, pulmonary macrophage, (or dust cell) is a type of macrophage, a professional phagocyte, found in the airways and at the level of the alveoli in the lungs, but separated from their walls. Activity of the alveolar macropha ...
s 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 splint ...
. 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 olde ...
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 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 ...
. For example significant changes can take place in COPD following infection with
rhinovirus The rhinovirus (from the grc, ῥίς, rhis "nose", , romanized: "of the nose", and the la, vīrus) is the most common viral infectious agent in humans and is the predominant cause of the common cold. Rhinovirus infection proliferates in tem ...
.
Fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
genera that are commonly found as
mycobiota Mycobiota (plural noun, no singular) are a group of all the fungi present in a particular geographic region (e.g. "the mycobiota of Ireland") or habitat type (e.g. "the mycobiota of cocoa"). Human mycobiota Mycobiota exist on the surface and in th ...
in the microbiota include '' Candida'', '' Malassezia'', '' Saccharomyces'', and ''
Aspergillus ' () is a genus consisting of several hundred mold species found in various climates worldwide. ''Aspergillus'' was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Mic ...
''.


Respiratory tract

The
lower respiratory tract The respiratory tract is the subdivision of the respiratory system involved with the process of Respiration (physiology), respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed ...
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 ...
, 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 the ...
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 Carina may refer to: Places Australia * Carina, Queensland, a suburb in Brisbane * Carina Heights, Queensland, a suburb in Brisbane * Carina, Victoria, a locality in Mildura Serbia * Carina, Osečina, a village in the Kolubara District ...
) into a right and left primary
bronchus 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 bronchu ...
. 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 bronchiole The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start o ...
s. These in turn supply air through alveolar ducts into the alveoli, where the exchange of gases take place. Oxygen breathed in, diffuses through the walls of the alveoli into the enveloping
capillaries A capillary is a small blood vessel from 5 to 10 micrometres (μm) in diameter. Capillaries are composed of only the tunica intima, consisting of a thin wall of simple squamous endothelial cells. They are the smallest blood vessels in the body: ...
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 The respiratory tract is the subdivision of the respiratory system involved with the process of respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed in through the nose to ...
are reinforced with hyaline cartilage in order to hold open the airways. The bronchioles have no cartilage and are surrounded instead by smooth muscle. Air is warmed to , humidified and cleansed by the conducting zone.
Particles In the physical sciences, a particle (or corpuscule in older texts) is a small localized object which can be described by several physical or chemical properties, such as volume, density, or mass. They vary greatly in size or quantity, from s ...
from the air being removed by the cilia on the
respiratory epithelium Respiratory epithelium, or airway epithelium, is a type of ciliated columnar epithelium found lining most of the respiratory tract as respiratory mucosa, where it serves to moisten and protect the airways. It is not present in the vocal cords o ...
lining the passageways, in a process called
mucociliary clearance Mucociliary clearance (MCC), mucociliary transport, or the mucociliary escalator, describes the self-clearing mechanism of the airways in the respiratory system. It is one of the two protective processes for the lungs in removing inhaled particl ...
.
Pulmonary stretch receptors Pulmonary stretch receptors are mechanoreceptors found in the lungs. When the lung expands, the receptors initiate the Hering-Breuer reflex, which reduces the respiratory rate. This signal is transmitted by vagus nerve. Increased firing from the ...
in the smooth muscle of the airways initiate a reflex known as the Hering–Breuer reflex 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 pulmonary circulation is a division of the circulatory system in all vertebrates. The circuit begins with deoxygenated blood returned from the body to the right atrium of the heart where it is pumped out from the right ventricle to the lungs ...
. The
bronchial circulation The bronchial circulation is the part of the circulatory system that supplies nutrients and oxygen to the cells that constitute the lungs, as well as carrying waste products away from them. It is complementary to the pulmonary circulation that brin ...
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 o ...
that leave the
aorta The aorta ( ) is the main and largest artery in the human body, originating from the left ventricle of the heart and extending down to the abdomen, where it splits into two smaller arteries (the common iliac arteries). The aorta distributes o ...
. 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 The pulmonary circulation is a division of the circulatory system in all vertebrates. The circuit begins with deoxygenated blood returned from the body to the right atrium of the heart where it is pumped out from the right ventricle to the lungs ...
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. Input from the parasympathetic nervous system 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, 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. 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 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 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 ...
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 the ...
,
bronchi 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 lungs that make up the respiratory tract, begin to form during the fourth week of embryogenesis 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 A branch is a part of a woody plant. Branch or branching may also refer to: Places ;Canada * Branch, Newfoundland and Labrador ;France * Branches, Yonne ;United States * Branch, Arkansas * Branch, Louisiana * Branch, Michigan * Branch, Mis ...
, 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 factor Fibroblast growth factors (FGF) are a family of cell signalling proteins produced by macrophages; they are involved in a wide variety of processes, most notably as crucial elements for normal development in animal cells. Any irregularities in their ...
s FGF10 and FGFR2b, and bone morphogenetic protein
BMP4 Bone morphogenetic protein 4 is a protein that in humans is encoded by ''BMP4'' gene. BMP4 is found on chromosome 14q22-q23. BMP4 is a member of the bone morphogenetic protein family which is part of the transforming growth factor-beta superfamil ...
. FGF10 is seen to have the most prominent role. FGF10 is a
paracrine signalling Paracrine signaling is a form of cell signaling, a type of cellular communication in which a cell produces a signal to induce changes in nearby cells, altering the behaviour of those cells. Signaling molecules known as paracrine factors diffuse over ...
molecule needed for epithelial branching, and SHH inhibits FGF10. The development of the alveoli is influenced by a different mechanism whereby continued bifurcation is stopped and the distal tips become dilated to form the alveoli. At the end of the fourth week the lung bud divides into two, the right and left primary bronchial buds on each side of the trachea. During the fifth week the right bud branches into three secondary bronchial buds and the left branches into two secondary bronchial buds. These give rise to the lobes of the lungs, three on the right and two on the left. Over the following week, the secondary buds branch into tertiary buds, about ten on each side. From the sixth week to the sixteenth week, the major elements of the lungs appear except the alveoli. From week 16 to week 26, the bronchi enlarge and lung tissue becomes highly vascularised. Bronchioles and alveolar ducts also develop. By week 26 the terminal bronchioles have formed which branch into two respiratory bronchioles. During the period covering the 26th week until birth the important
blood–air barrier The blood–air barrier or air–blood barrier, (alveolar–capillary barrier or membrane) exists in the gas exchanging region of the lungs. It exists to prevent air bubbles from forming in the blood, and from blood entering the alveoli. It is ...
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 i ...
, appear. The surfactant reduces the surface tension at the air-alveolar surface which allows expansion of the alveolar sacs. The alveolar sacs contain the primitive alveoli that form at the end of the alveolar ducts, and their appearance around the seventh month marks the point at which limited respiration would be possible, and the premature baby could survive.


Vitamin A deficiency

The developing lung is particularly vulnerable to changes in the levels of
vitamin A Vitamin A is a fat-soluble vitamin and an essential nutrient for humans. It is a group of organic compounds that includes retinol, retinal (also known as retinaldehyde), retinoic acid, and several provitamin A carotenoids (most notably ...
.
Vitamin A deficiency Vitamin A deficiency (VAD) or hypovitaminosis A is a lack of vitamin A in blood and tissues. It is common in poorer countries, especially among children and women of reproductive age, but is rarely seen in more developed countries. Nyctalopia (ni ...
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 Birth is the act or process of bearing or bringing forth offspring, also referred to in technical contexts as parturition. In mammals, the process is initiated by hormones which cause the muscular walls of the uterus to contract, expelling the f ...
, 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 The blood–air barrier or air–blood barrier, (alveolar–capillary barrier or membrane) exists in the gas exchanging region of the lungs. It exists to prevent air bubbles from forming in the blood, and from blood entering the alveoli. It is ...
. 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 The muscles of respiration are the muscles that contribute to inhalation and exhalation, by aiding in the expansion and contraction of the thoracic cavity. The diaphragm and, to a lesser extent, the intercostal muscles drive respiration during ...
, through the contraction of the diaphragm, and the
intercostal muscles Intercostal muscles are many different groups of muscles that run between the ribs, and help form and move the chest wall. The intercostal muscles are mainly involved in the mechanical aspect of breathing by helping expand and shrink the size of ...
which pull the
rib cage 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 sem ...
upwards as shown in the diagram. During breathing out the muscles relax, returning the lungs to their resting position. At this point the lungs contain the
functional residual capacity Functional residual capacity (FRC) is the volume of air present in the lungs at the end of passive expiration. At FRC, the opposing elastic recoil forces of the lungs and chest wall are in equilibrium and there is no exertion by the diaphragm ...
(FRC) of air, which, in the adult human, has a volume of about 2.5–3.0 litres. During heavy breathing as in
exertion Exertion is the physical or perceived use of energy.Newton's Third Law, Elert, Glenn. “Forces.” ''Viscosity – The Physics Hypertextbook'', physics.info/newton-first/. Exertion traditionally connotes a strenuous or costly ''effort'', resulting ...
, 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 Lung volumes and lung capacities refer to the volume of air in the lungs at different phases of the respiratory cycle. The average total lung capacity of an adult human male is about 6 litres of air. Tidal breathing is normal, resting breath ...
and capacities.


Protection

The lungs possess several characteristics which protect against infection. The respiratory tract is lined by
respiratory epithelium Respiratory epithelium, or airway epithelium, is a type of ciliated columnar epithelium found lining most of the respiratory tract as respiratory mucosa, where it serves to moisten and protect the airways. It is not present in the vocal cords o ...
or respiratory mucosa, with hair-like projections called cilia 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 Mucociliary clearance (MCC), mucociliary transport, or the mucociliary escalator, describes the self-clearing mechanism of the airways in the respiratory system. It is one of the two protective processes for the lungs in removing inhaled particl ...
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 Immunoglobulin A (Ig A, also referred to as sIgA in its secretory form) is an antibody that plays a role in the immune function of mucous membranes. The amount of IgA produced in association with mucosal membranes is greater than all other typ ...
which protects against respiratory infections; goblet cells secrete mucus which also contains several antimicrobial compounds such as defensins, antiproteases, and antioxidants. 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 macrophages, immune cells which engulf and destroy debris and microbes that enter the lung in a process known as
phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...
; and dendritic cells 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 and behind the mouth in the
oropharynx 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 struc ...
, and larger particles are trapped in
nasal hair Nasal hair or nose hair, is the hair in the human nose. Adult humans have hair in the nostrils. Nasal hair functions include filtering foreign particles from entering the nasal cavity, and collecting moisture. In support of the first function, the ...
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 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 (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 a ...
to angiotensin II. The lungs are involved in the blood's
acid–base homeostasis Acid–base homeostasis is the homeostatic regulation of the pH of the body's extracellular fluid (ECF). The proper balance between the acids and bases (i.e. the pH) in the ECF is crucial for the normal physiology of the body—and for cellu ...
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 trans ...
when breathing. The lungs also serve a protective role. Several blood-borne substances, such as a few types of prostaglandins,
leukotrienes Leukotrienes are a family of eicosanoid inflammatory mediators produced in leukocytes by the oxidation of arachidonic acid (AA) and the essential fatty acid eicosapentaenoic acid (EPA) by the enzyme arachidonate 5-lipoxygenase. Leukotri ...
, serotonin 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 and ...
, 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 oxygenat ...
s and prevent them from entering
arteries An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the pu ...
and causing strokes. The lungs also play a pivotal role in speech by providing air and airflow for the creation of vocal sounds, and other
paralanguage 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 relati ...
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 and gasps. 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, SFTPB and SFTPC, and napsin, expressed in type II pneumocytes. Other proteins with elevated expression in the lung are the dynein protein
DNAH5 Dynein axonemal heavy chain 5 is a protein that in humans is encoded by the ''DNAH5'' gene. ''DNAH5'' is a protein-coding gene.1 It provides the instructions for synthesizing a protein that belongs to a microtubule-associated protein complex made ...
in ciliated cells, and the secreted
SCGB1A1 Uteroglobin, or blastokinin, also known as secretoglobin family 1A member 1 (SCGB1A1), is a protein that in humans is encoded by the ''SCGB1A1'' gene. SCGB1A1 is the founding member of the secretoglobin family of small, secreted, disulfide-bridged ...
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 disease Respiratory diseases, or lung diseases, are pathological conditions affecting the organs and tissues that make gas exchange difficult in air-breathing animals. They include conditions of the respiratory tract including the trachea, bronchi, bro ...
s 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 ...
.
Cardiothoracic surgery Cardiothoracic surgery is the field of medicine involved in surgical treatment of organs inside the thoracic cavity — generally treatment of conditions of the heart (heart disease), lungs ( lung disease), and other pleural or mediastinal str ...
deals with surgery of the lungs including
lung volume reduction surgery Cardiothoracic surgery is the field of medicine involved in surgical treatment of organs inside the thoracic cavity — generally treatment of conditions of the heart (heart disease), lungs (lung disease), and other pleural or mediastinal struc ...
,
lobectomy Lobectomy means ''surgical excision of a lobe''. This may refer to a lobe of the lung (also simply called a lobectomy), a lobe of the thyroid ( hemithyroidectomy), a lobe of the brain (as in anterior temporal lobectomy), or a lobe of the liver ...
, pneumectomy 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 severi ...
, of the respiratory tract are
bronchitis Bronchitis is inflammation of the bronchi (large and medium-sized airways) in the lungs that causes coughing. Bronchitis usually begins as an infection in the nose, ears, throat, or sinuses. The infection then makes its way down to the bronchi. ...
and
bronchiolitis Bronchiolitis is inflammation of the small airways in the lungs. Acute bronchiolitis is due to a viral infection usually affecting children younger than two years of age. Symptoms may include fever, cough, runny nose, wheezing, and breathing ...
, and of the
pleurae The pulmonary pleurae (''sing.'' pleura) are the two opposing layers of serous membrane overlying the lungs and the inside of the surrounding chest walls. The inner pleura, called the visceral pleura, covers the surface of each lung and dips be ...
surrounding the lungs
pleurisy Pleurisy, also known as pleuritis, is inflammation of the membranes that surround the lungs and line the chest cavity ( pleurae). This can result in a sharp chest pain while breathing. Occasionally the pain may be a constant dull ache. Other sy ...
. 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 dis ...
s due to
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
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 Pneumonitis describes general inflammation of lung tissue. Possible causative agents include radiation therapy of the chest, exposure to medications used during chemo-therapy, the inhalation of debris (e.g., animal dander), aspiration, herbicide ...
. One major cause of
bacterial pneumonia Bacterial pneumonia is a type of pneumonia caused by bacterial infection. Types Gram-positive '' Streptococcus pneumoniae'' () is the most common bacterial cause of pneumonia in all age groups except newborn infants. ''Streptococcus 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, i ...
. Chronic infections often occur in those with immunodeficiency and can include a fungal infection by '' Aspergillus fumigatus'' that can lead to an
aspergilloma An aspergilloma is a clump of mold which exists in a body cavity such as a paranasal sinus or an organ such as the lung. By definition, it is caused by fungi of the genus ''Aspergillus''. Signs and symptoms People with aspergillomata typically rem ...
forming in the lung. Alcohol affects the lungs and can cause inflammatory alcoholic lung disease. Acute exposure to alcohol stimulates the beating of cilia in the respiratory epithelium. However, chronic exposure has the effect of desensitising the ciliary response which reduces
mucociliary clearance Mucociliary clearance (MCC), mucociliary transport, or the mucociliary escalator, describes the self-clearing mechanism of the airways in the respiratory system. It is one of the two protective processes for the lungs in removing inhaled particl ...
(MCC). MCC is an innate defense system protecting against pollutants and pathogens, and when this is disrupted the numbers of
alveolar macrophage An alveolar macrophage, pulmonary macrophage, (or dust cell) is a type of macrophage, a professional phagocyte, found in the airways and at the level of the alveoli in the lungs, but separated from their walls. Activity of the alveolar macropha ...
s 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 autocrin ...
s. Another consequence is the susceptibility to infection.


Blood-supply changes

A
pulmonary embolism Pulmonary embolism (PE) is a blockage of an artery in the lungs by a substance that has moved from elsewhere in the body through the bloodstream ( embolism). Symptoms of a PE may include shortness of breath, chest pain particularly upon breathin ...
is a blood clot that becomes lodged in the
pulmonary arteries A pulmonary artery is an artery in the pulmonary circulation that carries deoxygenated blood from the right side of the heart to the lungs. The largest pulmonary artery is the ''main pulmonary artery'' or ''pulmonary trunk'' from the heart, and ...
. The majority of emboli arise because of
deep vein thrombosis Deep vein thrombosis (DVT) is a type of venous thrombosis involving the formation of a blood clot in a deep vein, most commonly in the legs or pelvis. A minority of DVTs occur in the arms. Symptoms can include pain, swelling, redness, and enla ...
in the legs. Pulmonary emboli may be investigated using a
ventilation/perfusion scan A ventilation/perfusion lung scan, also called a V/Q lung scan, or ventilation/perfusion scintigraphy, is a type of medical imaging using scintigraphy and medical isotopes to evaluate the circulation of air and blood within a patient's lungs, in ...
, 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 A pulmonary artery is an artery in the pulmonary circulation that carries deoxygenated blood from the right side of the heart to the lungs. The largest pulmonary artery is the ''main pulmonary artery'' or ''pulmonary trunk'' from the heart, and ...
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 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 A pleural effusion is accumulation of excessive fluid in the pleural space, the potential space that surrounds each lung. Under normal conditions, pleural fluid is secreted by the parietal pleural capillaries at a rate of 0.6 millilitre per k ...
, or other substances such as air (
pneumothorax A pneumothorax is an abnormal collection of air in the pleural space between the lung and the chest wall. Symptoms typically include sudden onset of sharp, one-sided chest pain and shortness of breath. In a minority of cases, a one-way valve i ...
), blood (
hemothorax A hemothorax (derived from hemo- lood+ thorax hest plural ''hemothoraces'') is an accumulation of blood within the pleural cavity. The symptoms of a hemothorax may include chest pain and difficulty breathing, while the clinical signs may incl ...
), or rarer causes. These may be investigated using a chest X-ray or CT scan, and may require the insertion of a surgical drain until the underlying cause is identified and treated.


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, 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 Bronchitis is inflammation of the bronchi (large and medium-sized airways) in the lungs that causes coughing. Bronchitis usually begins as an infection in the nose, ears, throat, or sinuses. The infection then makes its way down to the bronchi. ...
, and emphysema, are all
obstructive lung disease Obstructive lung disease is a category of respiratory disease characterized by airway obstruction. Many obstructive diseases of the lung result from narrowing (obstruction) of the smaller bronchi and larger bronchioles, often because of excessiv ...
s 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 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), with symptom control such as
bronchodilator 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 ...
s, and with suppression of inflammation (such as through corticosteroids) in severe cases. A common cause of chronic bronchitis, and emphysema, is smoking; and common causes of bronchiectasis 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 In molecular biology, elastase is an enzyme from the class of ''proteases (peptidases)'' that break down proteins. In particular, it is a serine protease. Forms and classification Eight human genes exist for elastase: Some bacteria (includin ...
breaks down the elastin 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-p ...
,
alpha-1 antitrypsin Alpha-1 antitrypsin or α1-antitrypsin (A1AT, α1AT, A1A, or AAT) is a protein belonging to the serpin superfamily. It is encoded in humans by the ''SERPINA1'' gene. A protease inhibitor, it is also known as alpha1–proteinase inhibitor (A1PI ...
, and when there is a deficiency in this, emphysema can develop. With persistent stress from smoking, the
airway basal cell Airway basal cells are found deep in the respiratory epithelium, attached to, and lining the basement membrane. Basal cells are the stem cells or progenitors of the airway epithelium and can differentiate to replenish all of the epithelial cells ...
s 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 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 ...
, 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 oxygenat ...
, because of a restriction in the amount of lung tissue involved in respiration. These include
pulmonary fibrosis Pulmonary fibrosis is a condition in which the lungs become scarred over time. Symptoms include shortness of breath, a dry cough, feeling tired, weight loss, and nail clubbing. Complications may include pulmonary hypertension, respiratory failu ...
which can occur when the lung is inflamed for a long period of time.
Fibrosis Fibrosis, also known as fibrotic scarring, is a pathological wound healing in which connective tissue replaces normal parenchymal tissue to the extent that it goes unchecked, leading to considerable tissue remodelling and the formation of perma ...
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 disease Occupational lung diseases are work-related, lung conditions that have been caused or made worse by the materials a person is exposed to within the workplace. It includes a broad group of diseases, including occupational asthma, industrial bronch ...
s 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 ...
, autoimmune diseases 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, malign ...
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, then ...
from another part of the body. There are two main types of primary tumour described as either small-cell or
non-small-cell lung carcinoma Non-small-cell lung cancer (NSCLC) is any type of epithelial lung cancer other than small-cell lung carcinoma (SCLC). NSCLC accounts for about 85% of all lung cancers. As a class, NSCLCs are relatively insensitive to chemotherapy, compared to s ...
s. The major risk factor for cancer is smoking. Once a cancer is identified it is staged using scans such as a CT scan and a sample of tissue 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,
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. Chemothe ...
or a combination, or with the aim of symptom control. Lung cancer screening 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 ca ...
s include cystic fibrosis, pulmonary hypoplasia (an incomplete development of the lungs)
congenital diaphragmatic hernia Congenital diaphragmatic hernia (CDH) is a birth defect of the diaphragm. The most common type of CDH is a Bochdalek hernia; other types include Morgagni hernia, diaphragm eventration and central tendon defects of the diaphragm. Malformation of ...
, 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 In human anatomy, an azygos lobe is a normal anatomical variation of the upper lobe of the right lung. It is seen in 0.3% of the population. Embryologically, it arises from an anomalous lateral course of the azygos vein, in a pleural septum withi ...
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 A pneumothorax is an abnormal collection of air in the pleural space between the lung and the chest wall. Symptoms typically include sudden onset of sharp, one-sided chest pain and shortness of breath. In a minority of cases, a one-way valve i ...
(collapsed lung) is an abnormal collection of air in the
pleural space The pleural cavity, pleural space, or interpleural space is the potential space between the pleurae of the pleural sac that surrounds each lung. A small amount of serous pleural fluid is maintained in the pleural cavity to enable lubrication bet ...
that causes an uncoupling of the lung from the chest wall. The lung cannot expand against the air pressure inside the pleural space. An easy to understand example is a traumatic pneumothorax, where air enters the pleural space from outside the body, as occurs with puncture to the chest wall. Similarly, scuba divers ascending while holding their breath with their lungs fully inflated can cause air sacs ( alveoli) to burst and leak high pressure air into the pleural space.


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 cough is a sudden expulsion of air through the large breathing passages that can help clear them of fluids, irritants, foreign particles and microbes. As a protective reflex, coughing can be repetitive with the cough reflex following three ph ...
, a lung examination may be carried out. This exam includes
palpation Palpation is the process of using one's hands to check the body, especially while perceiving/diagnosing a disease or illness. Usually performed by a health care practitioner, it is the process of feeling an object in or on the body to determine ...
and auscultation. The areas of the lungs that can be listened to using a stethoscope are called the lung fields, 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
axilla The axilla (also, armpit, underarm or oxter) is the area on the human body directly under the shoulder joint. It includes the axillary space, an anatomical space within the shoulder girdle between the arm and the thoracic cage, bounded superior ...
e, 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 Muscle, musculature of the Human back, back, situated along the medial border of the scapula which allows for improved auscultation, listening to the lungs. Boundaries It has the followi ...
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 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 ...
.


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 Tidal volume (symbol VT or TV) is the volume of air moved into or out of the lungs during a normal breath. In a healthy, young human adult, tidal volume is approximately 500 ml per inspiration or 7 ml/kg of body mass. Mechanical vent ...
(normally 500-750mL); the
inspiratory reserve volume Lung volumes and lung capacities refer to the volume of air in the lungs at different phases of the respiratory cycle. The average total lung capacity of an adult human male is about 6 litres of air. Tidal breathing is normal, resting breath ...
and
expiratory reserve volume Lung volumes and lung capacities refer to the volume of air in the lungs at different phases of the respiratory cycle. The average total lung capacity of an adult human male is about 6 litres of air. Tidal breathing is normal, resting breath ...
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 Vital capacity (VC) is the maximum amount of air a person can inhale after a maximum exhalation. It is equal to the sum of inspiratory reserve volume, tidal volume, and expiratory reserve volume. It is approximately equal to Forced 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 Lung volumes and lung capacities refer to the volume of air in the lungs at different phases of the respiratory cycle. The average total lung capacity of an adult human male is about 6 litres of air. Tidal breathing is normal, resting breath ...
. Pulmonary
plethysmograph A plethysmograph is an instrument for measuring changes in volume within an organ or whole body (usually resulting from fluctuations in the amount of blood or air it contains). The word is derived from the Greek "plethysmos" (increasing, enla ...
s are used to measure
functional residual capacity Functional residual capacity (FRC) is the volume of air present in the lungs at the end of passive expiration. At FRC, the opposing elastic recoil forces of the lungs and chest wall are in equilibrium and there is no exertion by the diaphragm ...
. 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 Vital capacity (VC) is the maximum amount of air a person can inhale after a maximum exhalation. It is equal to the sum of inspiratory reserve volume, tidal volume, and expiratory reserve volume. It is approximately equal to Forced Vital Capacity ( ...
. In particular, how much a person is able to exhale in one second (called forced expiratory volume (FEV1)) as a proportion of how much they are able to exhale in total (FEV). This ratio, the FEV1/FEV ratio, is important to distinguish whether a lung disease is restrictive or obstructive. Another test is that of the lung's
diffusing capacity Diffusing capacity of the lung (DL) (also known as Transfer factor is another expression for the formerly used diffusing capacity.) measures the transfer of gas from air in the lung, to the red blood cells in lung blood vessels. It is part of a c ...
– 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 ...
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
tuatara Tuatara (''Sphenodon punctatus'') are reptiles endemic to New Zealand. Despite their close resemblance to lizards, they are part of a distinct lineage, the order Rhynchocephalia. The name ''tuatara'' is derived from the Māori language and m ...
s,
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 j ...
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 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 living ...
and
monitor lizards Monitor lizards are lizards in the genus ''Varanus,'' the only extant genus in the family Varanidae. They are native to Africa, Asia, and Oceania, and one species is also found in the Americas as an invasive species. About 80 species are recogn ...
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, and azhdarchoids, the hindlimbs. Reptilian lungs typically receive air via expansion and contraction of the ribs driven by axial muscles and buccal pumping. Crocodilians also rely on the
hepatic The liver is a major organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the synthesis of proteins and biochemicals necessary for digestion and growth. In humans, it is ...
piston method, in which the liver is pulled back by a muscle anchored to the
pubic bone In vertebrates, the pubic region ( la, pubis) is the most forward-facing ( ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior ...
(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 Boyle's law, also referred to as the Boyle–Mariotte law, or Mariotte's law (especially in France), is an experimental gas law that describes the relationship between pressure and volume of a confined gas. Boyle's law has been stated as: The ...
.
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 The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of ...
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 carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
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 Positive pressure is a pressure within a system that is greater than the environment that surrounds that system. Consequently, if there is any leak from the positively pressured system it will egress into the surrounding environment. This is in ...
system to get air to their lungs, forcing air down into the lungs by
buccal pumping Buccal pumping is "breathing with one's cheeks": a method of ventilation used in respiration in which the animal moves the floor of its mouth in a rhythmic manner that is externally apparent.Brainerd, E. L. (1999). New perspectives on the evolutio ...
. This is distinct from most higher vertebrates, who use a breathing system driven by
negative pressure Negative pressure may refer to: * Negative value of a pressure variable * Negative room pressure, a ventilation technique used to avoid contaminating outside areas * Negative pressure ventilator, also known as an iron lung * Negative-pressure wou ...
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 salamander Plethodontidae, or lungless salamanders, are a family of salamanders. Most species are native to the Western Hemisphere, from British Columbia to Brazil, although a few species are found in Sardinia, Europe south of the Alps, and South Korea. In ...
s, 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 The Bornean flat-headed frog (''Barbourula kalimantanensis'') is a species of frog in the family Bombinatoridae. Although many salamanders are lungless, the Bornean flat-headed frog is the first frog known to have no lungs. Djoko Iskandar, an In ...
and '' Atretochoana eiselti'', a
caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics o ...
. 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 The Australian lungfish (''Neoceratodus forsteri''), also known as the Queensland lungfish, Burnett salmon and barramunda, is the only surviving member of the family Neoceratodontidae. It is one of only six extant lungfish species in the world. ...
, there is only a single lung, albeit divided into two lobes. Other lungfish and ''
Polypterus ''Polypterus'' is a genus of freshwater fish in the bichir family (Polypteridae) of order Polypteriformes. The type species is the Nile bichir (''P. bichir''). Fish in this genus live in various areas in Africa. ''Polypterus'' is the only kno ...
'', 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 ...
. 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 chordate ...
s have lung-like structures that serve a similar respiratory purpose as, but are not evolutionarily related to, vertebrate lungs. Some
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...
s, 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 lungs used for atmospheric gas exchange. Some species of spider have four pairs of book lungs but most have two pairs. Scorpions have spiracles on their body for the entrance of air to the book lungs. The
coconut crab 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 lung A branchiostegal lung is a respiration organ used by some air-breathing arthropods. It is one of the most significant adaptations of some crabs and hermit crabs such as the coconut crab to their terrestrial habitats. The branchiostegal (gill) tiss ...
s 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 The mantle (also known by the Latin word pallium meaning mantle, robe or cloak, adjective pallial) is a significant part of the anatomy of molluscs: it is the dorsal body wall which covers the visceral mass and usually protrudes in the form of ...
. An externally located opening called the
pneumostome The pneumostome or breathing pore is a respiratory opening of the external body anatomy of an air-breathing land slug or land snail. It is a part of the respiratory system of gastropods. It is an opening in the right side of the mantle of a ...
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 c ...
s and the gas bladders of today's
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
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 ...
, that allowed early fish to gulp air under oxygen-poor conditions. These outpocketings first arose in the bony fish. In most of the
ray-finned fish 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,
trout Trout are species of freshwater fish belonging to the genera '' Oncorhynchus'', '' Salmo'' and '' Salvelinus'', all of the subfamily Salmoninae of the family Salmonidae. The word ''trout'' is also used as part of the name of some non-salm ...
, 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
eel Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
s have retained the physostome condition with the sac being open to the esophagus. In more basal bony fish, such as the
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
,
bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pu ...
, bowfin and the
lobe-finned fish Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includ ...
, 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 are ...
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 Drowning is a type of suffocation induced by the submersion of the mouth and nose in a liquid. Most instances of fatal drowning occur alone or in situations where others present are either unaware of the victim's situation or unable to offer as ...
*
Interstitial lung disease Interstitial lung disease (ILD), or diffuse parenchymal lung disease (DPLD), is a group of respiratory diseases affecting the interstitium (the tissue and space around the alveoli (air sacs)) of the lungs. It concerns alveolar epithelium, pu ...
* Liquid breathing *
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 ma ...
* Lung-on-a-chip * Secarecytosis *
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