Anatomy (from the Greek ἀνατέμνω – anatemnō, "I cut up, cut open", from ἀνά – ana, "on, upon", and τέμνω – temnō, "I cut") is the study of the body plan of animals. In some of its facets, anatomy is closely related to embryology, comparative anatomy and comparative embryology, through common roots in evolution. Human anatomy is important in medicine.
The discipline of anatomy is subdivided into gross or macroscopic anatomy, and microscopic anatomy. Gross anatomy is the study of structures that can, when suitably presented or dissected, be seen by unaided vision with the naked eye. Microscopic anatomy is the study of structures on a microscopic scale, including histology (the study of tissues) and cytology (the study of cells).
The history of anatomy is characterized by a continual development in understanding of the functions of the organs and structures of the human body. Methods have also improved dramatically, advancing from examination of animals through dissection of cadavers (dead human bodies) to 20th century techniques including X-ray, ultrasound, and magnetic resonance imaging.
Anatomy is the scientific study of the structure of living things including their systems, organs, and tissues. It includes the appearance and position of the various parts, the materials from which they are composed, their locations and their relationships with other parts. Anatomy is quite distinct from physiology and biochemistry which deals with the functions of those parts. For example, an anatomist is concerned with the shape, size, position, structure, blood supply and enervation of an organ such as the liver while a physiologist will be interested in the production of bile and the role of the liver in nutrition and regulating body functions.
The discipline of anatomy can be subdivided into a number of branches including gross or macroscopic anatomy and microscopic anatomy. Gross anatomy is the study of structures that are large enough to be seen with the naked eye. It can be studied using both invasive and non-invasive methods with the goal of obtaining information about the structure and organization of organs and systems. Methods used include dissection, in which the body is surgically opened and its organs studied, and endoscopy, in which a video camera-equipped instrument is inserted through a small incision in the body wall and used to explore the internal organs and other structures. Angiography is a proceeding in which blood vessels are visualized after being injected with an opaque dye, X-rays and magnetic resonance imaging being used for this purpose. Microscopic anatomy is the study of structures on a microscopic scale, including histology (the study of tissues), and embryology (the study of the human organism in its immature condition).
The term "anatomy" is commonly taken to refer to human anatomy. However, substantially the same structures and tissues are found throughout the animal kingdom and the term also includes the anatomy of other animals. The structure and tissues of plants are of a dissimilar nature and they are studied in plant anatomy.
The kingdom Animalia contains multicellular organisms that are heterotrophic and motile (although some have secondarily adopted a sessile lifestyle). Most animals have bodies differentiated into separate tissues and these animals are known as metazoans or eumetazoans. They have an internal digestive chamber, with one or two openings, the gametes are produced in multicellular sex organs and the zygotes include a blastula stage in their embryonic development. Metazoans do not include the sponges, phylum Porifera, or the Placozoans which have undifferentiated cells.
Animal cells do not have a cell wall and do not contain chloroplasts. If a vacuole is present, it is smaller than that found in a plant cell. The body tissues are composed of a great variety of cells such as muscle, nerve and blood cells. Each typically has a cell membrane formed of phospholipids, cytoplasm and a nucleus. Animal tissues can be grouped into four basic types: connective, epithelial, muscle and nervous. Even the simplest invertebrates contain at least two types and body structures and organs of higher animals are formed from multiple tissue types.
Connective tissues are fibrous tissues and are made up of cells scattered among non-living material which is called the extracellular matrix. Connective tissue gives shape to organs and holds them in place. The main types are loose connective tissue, adipose tissue, fibrous connective tissue, blood, cartilage and bone. The extracellular matrix contains proteins, one of the commonest of which is collagen, and plays an important part in organizing and maintaining tissues. The matrix can be modified to form a skeleton to support or protect the body. An exoskeleton is a thickened, rigid cuticle which is stiffened by mineralization, as in crustaceans or by the cross-linking of its proteins as in insects. An endoskeleton is internal and is present in all higher animals and many lower ones.
Epithelial tissue is composed of closely packed cells with little intercellular space. They can be squamous (flat), cuboidal or columnar and rest on a basal lamina, a membrane composed of extracellular matrix secreted by the epithelial cells. The cells are bound closely to each other by cell adhesion molecules. One or more layers of epithelial cells line the cavities and surfaces of structures throughout the body including the external surface, the respiratory surface and the gut. The epithelial cells on the external surface of the body typically secrete an extracellular matrix in the form of a cuticle. In simple animals this may just be a coat of glycoproteins. In more advanced animals, many glands are formed of epithelial cells.
Muscle cells (myocytes) form the active contractile tissue of the body. Muscle tissue functions to produce force and cause motion, either locomotion or movement within internal organs. Muscle is formed of contractile filaments and is separated into three types; smooth muscle, skeletal muscle and obliquely striated muscle. Smooth muscle has no striations when examined microscopically. It contracts slowly but maintains contractability over a wide range of stretch lengths. It is found in such organs as sea anemone tentacles and the body wall of sea cucumbers. Cross-striated muscle contracts rapidly but has a limited range of extension. It is found in the movement of appendages and jaws. Obliquely striated muscle is intermediate between the other two. The filaments are staggered and this is the type of muscle found in earthworms that can extend slowly or make rapid contractions. In higher animals striated muscles occur in bundles attached to bone to provide movement and are often arranged in antagonistic sets. Smooth muscle is found in the inner linings of organs and cardiac muscle is found in the heart, allowing it to contract and pump blood round the body.
Nervous tissue is composed of many nerve cells known as neurons which transmit information. In radially symmetrical animals such as sea anemones and jellyfish the nerves form a nerve net but in bilaterally symmetrical animals they are organized longitudinally into bundles. In simple animals, receptor neurons in the body wall cause a local reaction to a stimulus. In more complex animals, specialised receptor cells such as chemoreceptors and photoreceptors are found in groups and send messages along neurons to other parts of the organism. Neurons can be connected together in ganglia. In higher animals these form the sense organs and there is a central nervous system (brain and spinal cord) and a peripheral nervous system. The latter consists of sensory nerves that transmit information from sense organs and motor nerves that stimulate muscles to contract.
Human anatomy, including gross human anatomy and histology, is primarily the scientific study of the morphology of the adult human body. It differs from physiology in that anatomy is only the structures involved, and Physiology is the way those structures actually work.
Generally, students of certain biological sciences, paramedics, prosthetists and orthotists, physiotherapists, occupational therapists, nurses, and medical students learn gross anatomy and microscopic anatomy from anatomical models, skeletons, textbooks, diagrams, photographs, lectures and tutorials, and in addition, medical students generally also learn gross anatomy through practical experience of dissection and inspection of cadavers. The study of microscopic anatomy (or histology) can be aided by practical experience examining histological preparations (or slides) under a microscope. 
Human anatomy, physiology and biochemistry are complementary basic medical sciences, which are generally taught to medical students in their first year at medical school. Human anatomy can be taught regionally or systemically; that is, respectively, studying anatomy by bodily regions such as the head and chest, or studying by specific systems, such as the nervous or respiratory systems. The major anatomy textbook, Gray's Anatomy, has been reorganized from a systems format to a regional format, in line with modern teaching methods. A thorough working knowledge of anatomy is required by physicians, especially surgeons and doctors working in some diagnostic specialties, such as histopathology and radiology. 
Academic human anatomists are usually employed by universities, medical schools or teaching hospitals. They are often involved in teaching anatomy, and research into certain systems, organs, tissues or cells.
The major systems of the human body are the:
All vertebrates are built along the basic chordate body plan: a stiff rod running through the length of the animal (vertebral column or notochord), with a hollow tube of nervous tissue, the spinal cord above it and the gastrointestinal tract below. Nervous tissue is derived from the ectoderm, connective tissues are derived from mesoderm, and gut is derived from the endoderm. At the posterior end is a tail which continues the spinal cord and vertebrae but not the gut. The mouth is found at the anterior end of the animal, and the anus at the base of the tail. The defining characteristic of a vertebrate is the vertebral column, in which the notochord has been replaced by a segmented series of stiffer elements (vertebrae) separated by mobile joints (intervertebral discs), which each contain a soft central core (nucleus pulposus), derived from the notochord of the embryo. However, a few vertebrates have secondarily lost this anatomy, retaining the notochord into adulthood. These include the sturgeon and the coelacanth. Jawed vertebrates are typified by paired appendages, fins or legs, which may be secondarily lost. The limbs of vertebrates are considered to be homologous because the same underlying skeletal structure was inherited from their last common ancestor. This is one of the arguments put forward by Charles Darwin to support his theory of evolution.
The body of a fish is divided into a head, trunk and tail, although the divisions between the three are not always externally visible. The skeleton, which forms the support structure inside the fish, is either made of cartilage (cartilaginous fish) or bone (bony fishes). The main skeletal element is the vertebral column, composed of articulating vertebrae which are lightweight yet strong. The ribs attach to the spine and there are no limbs or limb girdles. The main external features of the fish, the fins, are composed of bony spines and soft rays and, with the exception of the caudal fins, have no direct connection with the spine. They are supported by the muscles which compose the main part of the trunk. The heart has two chambers and pumps the blood through the respiratory surfaces of the gills and on round the body in a single circulatory loop. The eyes are adapted for seeing underwater and have only local vision. There is an inner ear but no external or middle ear. Low frequency vibrations are detected by the lateral line system on the flanks which responds to nearby movements and to changes in water pressure.
Sharks and rays are basal fish with numerous primitive anatomical features similar to those of ancient fish, including skeletons composed of cartilage. Their bodies tend to be dorso-ventrally flattened, they usually have five pairs of gill slits and a large mouth set on the underside of the head. The dermis is covered with separate dermal placoid scales. They have a cloaca into which the urinary and genital passages open, but not a swim bladder. Cartilaginous fish produce a small number of large, yolky eggs. Some species are ovoviviparous and the young develop internally but others are oviparous and the larvae develop externally in egg cases.
The bony fish lineage shows more derived anatomical traits, often with major evolutionary changes from the features of ancient fish. They have a bony skeleton, are generally laterally flattened, have five pairs of gills protected by an operculum, and a mouth at or near the tip of the snout. The dermis is covered with overlapping scales. Bony fish have a swim bladder which helps them maintain a constant depth in the water column, but not a cloaca. They mostly spawn a large number of small eggs with little yolk which they broadcast into the water column.
Amphibians are a class of animals comprising frogs, salamanders and caecilians. They are tetrapods, but the caecilians and a few species of salamander have either no limbs or their limbs are much reduced in size. Their main bones are hollow and lightweight and are fully ossified and the vertebrae interlock with each other and have articular processes. Their ribs are usually short and may be fused to the vertebrae. Their skulls are mostly broad and short, and are often incompletely ossified. Their skin contains little keratin and lacks scales, but contains many mucous glands and in some species, poison glands. The hearts of amphibians have three chambers, two atria and one ventricle. They have a urinary bladder and nitrogenous waste products are excreted primarily as urea. Amphibians breathe by means of a pump action in which air is first drawn into the buccopharyngeal region through the nostrils. These are then closed and the air is forced into the lungs by contraction of the throat. They supplement this with gas exchange through the skin which needs to be kept moist.
In frogs the pelvic girdle is robust and the hind legs are much longer and stronger than the forelimbs. The feet have four or five digits and the toes are often webbed for swimming or have suction pads for climbing. Frogs have large eyes and no tail. Salamanders resemble lizards in appearance; their short legs project sideways, the belly is close to or in contact with the ground and they have a long tail. Caecilians superficially resemble earthworms and are limbless. They burrow by means of zones of muscle contractions which move along the body and they swim by undulating their body from side to side.
Reptiles are a class of animals comprising turtles, tuataras, lizards, snakes and crocodiles. They are tetrapods, but the snakes and a few species of lizards either have no limbs or their limbs are much reduced in size. Their bones are better ossified and their skeletons stronger than those of amphibians. The teeth are conical and mostly uniform in size. The surface cells of the epidermis are modified into horny scales which create a waterproof layer. Reptiles are unable to use their skin for respiration as do amphibians and have a more efficient respiratory system drawing air into their lungs by expanding their chest walls. The heart resembles that of the amphibian but there is a septum which more completely separates the oxygenated and deoxygenated bloodstreams. The reproductive system is designed for internal fertilisation, with a copulatory organ present in most species. The eggs are surrounded by amniotic membranes which prevents them from drying out and are laid on land, or develop internally in some species. The bladder is small as nitrogenous waste is excreted as uric acid.
Turtles are notable for their protective shells. They have an inflexible trunk encased in a horny carapace above and a plastron below. These are formed from bony plates embedded in the dermis which are overlain by horny ones and are partially fused with the ribs and spine. The neck is long and flexible and the head and the legs can be drawn back inside the shell. Turtles are vegetarians and the typical reptile teeth have been replaced by sharp, horny plates. In aquatic species, the front legs are modified into flippers.
Tuataras superficially resemble lizards but the lineages diverged in the Triassic era. There is one living species, Sphenodon punctatus. The skull has two openings (fenestrae) on either side and the jaw is rigidly attached to the skull. There is one row of teeth in the lower jaw and this fits between the two rows in the upper jaw when the animal chews. The teeth are merely projections of bony material from the jaw and eventually wear down. The brain and heart are more primitive than is the case in other reptiles and the lungs have a single chamber and lack bronchi. The tuatara has a well-developed parietal eye on its forehead.
Lizards have skulls with only one fenestra on each side, the lower bar of bone below the second fenestra having been lost. This results in the jaws being less rigidly attached which allows the mouth to open wider. Lizards are mostly quadrupeds, with the trunk held off the ground by short, sideways-facing legs, but a few species have no limbs and resemble snakes. Lizards have moveable eyelids, eardrums are present and some species have a central parietal eye.
Snakes are closely related to lizards, having branched off from a common ancestral lineage during the Cretaceous era, and they share many of the same features. The skeleton consists of a skull, a hyoid bone, spine and ribs though a few species retain a vestige of the pelvis and rear limbs in the form of pelvic spurs. The bar under the second fenestra has also been lost and the jaws have extreme flexibility allowing the snake to swallow its prey whole. Snakes lack moveable eyelids, the eyes being covered by transparent "spectacle" scales. They do not have eardrums but can detect ground vibrations through the bones of their skull. Their forked tongues are used as organs of taste and smell and some species have sensory pits on their heads enabling them to locate warm-blooded prey.
Crocodilians are large, low-slung aquatic reptiles with long snouts and large numbers of teeth. The head and trunk are dorso-ventrally flattened and the tail is laterally compressed. It undulates from side to side to force the animal through the water when swimming. The tough keratinised scales provide body armour and some are fused to the skull. The nostrils, eyes and ears are elevated above the top of the flat head enabling them to remain above the surface of the water when the animal is floating. Valves seal the nostrils and ears when it is submerged. Unlike other reptiles, crocodilians have hearts with four chambers allowing complete separation of oxygenated and deoxygenated blood.
Birds are tetrapods but though their hind limbs are used for walking or hopping, their front limbs are wings covered with feathers and adapted for flight. Birds are endothermic, have a high metabolic rate, a light skeletal system and powerful muscles. The long bones are thin, hollow and very light. Air sac extensions from the lungs occupy the centre of some bones. The sternum is wide and usually has a keel and the caudal vertebrae are fused. There are no teeth and the narrow jaws are adapted into a horn-covered beak. The eyes are relatively large, particularly in nocturnal species such as owls. They face forwards in predators and sideways in ducks.
The feathers are outgrowths of the epidermis and are found in localized bands from where they fan out over the skin. Large flight feathers are found on the wings and tail, contour feathers cover the bird's surface and fine down occurs on young birds and under the contour feathers of water birds. The only cutaneous gland is the single uropygial gland near the base of the tail. This produces an oily secretion that waterproofs the feathers when the bird preens. There are scales on the legs and feet and claws on the tips of the toes.
Mammals are a diverse class of animals, mostly terrestrial but some are aquatic and others have evolved flapping or gliding flight. They mostly have four limbs but some aquatic mammals have no limbs or limbs modified into fins and the forelimbs of bats are modified into wings. The legs of most mammals are situated below the trunk, which is held well clear of the ground. The bones of mammals are well ossified and their teeth, which are usually differentiated, are coated in a layer of prismatic enamel. The teeth are shed once (milk teeth) during the animal's lifetime or not at all, as is the case in cetaceans. Mammals have three bones in the middle ear and a cochlea in the inner ear. They are clothed in hair and their skin contains glands which secrete sweat. Some of these glands are specialised as mammary glands, producing milk to feed the young. Mammals breathe with lungs and have a muscular diaphragm separating the thorax from the abdomen which helps them draw air into the lungs. The mammalian heart has four chambers and oxygenated and deoxygenated blood are kept entirely separate. Nitrogenous waste is excreted primarily as urea.
Mammals are amniotes, and most are viviparous, giving birth to live young. The exception to this are the egg-laying monotremes, the platypus and the echidnas of Australia. Most other mammals have a placenta through which the developing foetus obtains nourishment, but in marsupials, the foetal stage is very short and the immature young is born and finds its way to its mother's pouch where it latches on to a nipple and completes its development.
Invertebrates constitute a vast array of living organisms ranging from the simplest unicellular eukaryotes to such complex creatures as the octopus, lobster and dragonfly. By definition, none of them has a backbone. The cells of single-cell protozoans have the same basic structure as those of multicellular animals but some parts are specialised into the equivalent of tissues and organs. Locomotion is often provided by cilia or flagella or may proceed via the advance of pseudopodia, food may be gathered by phagocytosis, energy needs may be supplied by photosynthesis and the cell may be supported by an endoskeleton or an exoskeleton. Some protozoans can form multicellular colonies.
Metazoans are multicellular organism, different groups of cells of which have separate functions. The most basic types of metazoan tissues are epithelium and connective tissue, both of which are present in nearly all invertebrates. The outer surface of the epidermis is normally formed of epithelial cells and secretes an extracellular matrix which provides support to the organism. An endoskeleton derived from the mesoderm is present in echinoderms, sponges and some cephalopods. Exoskeletons are derived from the epidermis and is composed of chitin in arthropods (insects, spiders, ticks, shrimps, crabs, lobsters). Calcium carbonate constitutes the shells of molluscs, brachiopods and some tube-building polychaete worms and silica forms the exoskeleton of the microscopic diatoms and radiolaria. Other invertebrates may have no rigid structures but the epidermis may secrete a variety of surface coatings such as the pinacoderm of sponges, the gelatinous cuticle of cnidarians (polyps, sea anemones, jellyfish) and the collagenous cuticle of annelids. The outer epithelial layer may include cells of several types including sensory cells, gland cells and stinging cells. There may also be protrusions such as microvilli, cilia, bristles, spines and tubercles.
Insects possess segmented bodies supported by a hard-jointed outer covering made mostly of chitin. The segments of the body are organized into three distinct parts, a head, a thorax and an abdomen. The head typically bears a pair of sensory antennae, a pair of compound eyes, one to three simple eyes (ocelli) and three sets of modified appendages that form the mouthparts. The thorax has three pairs of segmented legs, one pair each for the three segments that compose the thorax and one or two pairs of wings. The abdomen is composed of eleven segments, some of which may be fused and houses the digestive, respiratory, excretory and reproductive systems. There is considerable variations between species and many adaptations to the body parts, especially wings, legs, antennae and mouthparts.
In 1600 BCE, the Edwin Smith Surgical Papyrus described the heart, its vessels, liver, spleen, kidneys, hypothalamus, uterus and bladder, and showed blood vessels from the heart. The Ebers Papyrus (c. 1550 BCE) features a "treatise on the heart", with vessels carrying all the body's fluids to or from every member of the body.
The anatomy of the muscles and skeleton is described in the Hippocratic Corpus, written by unknown authors. Aristotle described vertebrate anatomy based on animal dissection. Praxagoras identified the difference between arteries and veins. Also in the 4th century BCE, Herophilos and Erasistratus produced more accurate anatomical descriptions based on vivisection of criminals in Alexandria during the Ptolemaic dynasty.
In the 2nd century, Galen wrote the final and highly influential anatomy treatise of ancient times. He compiled existing knowledge and studied organs through vivisection of animals. Galen's drawings, based mostly on dog anatomy, became effectively the only anatomical textbook for the next thousand years. His work was known to Renaissance doctors only through Islamic Golden Age medicine until it was translated from the Greek some time in the 15th century.
Between 1275 and 1326, the anatomists Mondino de Luzzi, Alessandro Achillini and Antonio Benivieni at Bologna carried out the first systematic human dissections since ancient times. Mondino's Anatomy of 1316 was the first textbook in the mediaeval rediscovery of human anatomy. It describes the body in the order followed in Mondino's dissections, starting with the abdomen, then the thorax, then the head and limbs. It was the standard anatomy textbook for the next century.
Andreas Vesalius (1514–1564) (Latinized from Andries van Wezel), professor of anatomy at the University of Padua, is considered the founder of modern human anatomy. Originally from Brabant, Vesalius published the influential book De humani corporis fabrica ("the structure of the human body"), a large format book in seven volumes, in 1543. The accurate and intricately detailed illustrations, often in allegorical poses against Italianate landscapes, are thought to have been made by the artist Jan van Calcar, a pupil of Titian.
The artist Leonardo da Vinci (1452–1519) was trained in anatomy by Andrea del Verrocchio. He made use of his anatomical knowledge in his artwork, making many sketches of skeletal structures, muscles and organs of humans and other vertebrates which he dissected.
In England, anatomy was the subject of the first public lectures given in any science; these were given by the Company of Barbers and Surgeons in the 16th century, joined in 1583 by the Lumleian lectures in surgery at the Royal College of Physicians.
To obtain corpses for 19th century anatomy lessons, criminals known as 'Resurrectionists' resorted to body snatching, digging up newly-buried bodies from graveyards for sale to anatomists. Some graveyards were in consequence protected with watchtowers. The practice was halted in Britain by the Anatomy Act of 1832.
The teaching of anatomy in Britain was transformed by Sir John Struthers, Regius Professor of Anatomy at the University of Aberdeen from 1863 to 1889. He was responsible for setting up the system of three years of "pre-clinical" academic teaching in the sciences underlying medicine, including especially anatomy. This system lasted until the reform of medical training in 1993 and 2003. As well as teaching, he collected many vertebrate skeletons for his museum of Comparative anatomy, published over 70 research papers, and became famous for his public dissection of the Tay Whale.
During the 19th century, anatomical research was extended to include the histology and developmental biology of both humans and animals. Demand for bodies grew so great that body-snatching and even anatomy murder were practised to obtain cadavers. The British Parliament passed the Anatomy Act 1832, which finally provided for an adequate and legitimate supply of corpses by allowing dissection of destitutes. From 1822 the Royal College of Surgeons regulated the teaching of anatomy in medical schools. Medical museums provided examples in comparative anatomy, and were often used in teaching.
Anatomy in the 20th century took advantage of technological developments and growing understanding of sciences such as evolutionary and molecular biology to create a thorough understanding of the body's organs and structures. Techniques including X-ray, magnetic resonance imaging, computed tomography and ultrasound imaging enabled researchers and practitioners to examine organs, living or dead, in unprecedented detail.
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