Anatomy

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De humani corporis fabrica
16th century, marking the rebirth of anatomy

Anatomy (from

phylogeny,[3] as these are the processes by which anatomy is generated, both over immediate and long-term timescales. Anatomy and physiology, which study the structure and function of organisms and their parts respectively, make a natural pair of related disciplines, and are often studied together. Human anatomy is one of the essential basic sciences that are applied in medicine.[4]

Anatomy is a complex and dynamic field that is constantly evolving as new discoveries are made. In recent years, there has been a significant increase in the use of advanced imaging techniques, such as MRI and CT scans, which allow for more detailed and accurate visualizations of the body's structures.

The discipline of anatomy is divided into

superficial anatomy. Microscopic anatomy involves the use of optical instruments in the study of the tissues of various structures, known as histology, and also in the study of cells
.

The

Etymology and definition

A dissected body, lying prone on a table, by Charles Landseer

Derived from the Greek ἀνατομή anatomē "dissection" (from ἀνατέμνω anatémnō "I cut up, cut open" from ἀνά aná "up," and τέμνω témnō "I cut"),[7] anatomy is the scientific study of the structure of organisms including their systems, organs and tissues. It includes the appearance and position of the various parts, the materials from which they are composed, and their relationships with other parts. Anatomy is quite distinct from physiology and biochemistry, which deal respectively with the functions of those parts and the chemical processes involved. For example, an anatomist is concerned with the shape, size, position, structure, blood supply and innervation of an organ such as the liver; while a physiologist is interested in the production of bile, the role of the liver in nutrition and the regulation of bodily functions.[8]

The discipline of anatomy can be subdivided into a number of branches, including gross or

Microscopic anatomy is the study of structures on a microscopic scale, along with histology (the study of tissues), and embryology (the study of an organism in its immature condition).[3] Regional anatomy is the study of the interrelationships of all of the structures in a specific body region, such as the abdomen. In contrast, systemic anatomy is the study of the structures that make up a discrete body system—that is, a group of structures that work together to perform a unique body function, such as the digestive system.[10]

Anatomy 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.[3] Methods used include dissection, in which a body is 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 using X-rays or magnetic resonance angiography are methods to visualize blood vessels.[11][12][13][14]

The term "anatomy" is commonly taken to refer to

human anatomy. However, substantially similar structures and tissues are found throughout the rest of the animal kingdom, and the term also includes the anatomy of other animals. The term zootomy is also sometimes used to specifically refer to non-human animals. The structure and tissues of plants are of a dissimilar nature and they are studied in plant anatomy.[8]

Animal tissues

The

embryonic development. Metazoans do not include the sponges, which have undifferentiated cells.[15]

Unlike

animal cells have neither a cell wall nor chloroplasts. Vacuoles, when present, are more in number and much smaller than those in the plant cell. The body tissues are composed of numerous types of cells, including those found in muscles, nerves and skin. Each typically has a cell membrane formed of phospholipids, cytoplasm and a nucleus. All of the different cells of an animal are derived from the embryonic germ layers. Those simpler invertebrates which are formed from two germ layers of ectoderm and endoderm are called diploblastic and the more developed animals whose structures and organs are formed from three germ layers are called triploblastic.[16] All of a triploblastic animal's tissues and organs are derived from the three germ layers of the embryo, the ectoderm, mesoderm and endoderm
.

Animal tissues can be grouped into four basic types:

.

Hyaline cartilage at high magnification (H&E stain
)

Connective tissue

mineralization, as in crustaceans or by the cross-linking of its proteins as in insects. An endoskeleton is internal and present in all developed animals, as well as in many of those less developed.[16]

Epithelium

Gastric mucosa at low magnification (H&E stain
)

glycoproteins.[16] In more advanced animals, many glands are formed of epithelial cells.[20]

Muscle tissue

Cross section through skeletal muscle and a small nerve at high magnification (H&E stain
)

, allowing it to contract and pump blood round the body.

Nervous tissue

voluntary muscle, and the autonomic nervous system which involuntarily controls smooth muscle, certain glands and internal organs, including the stomach.[25]

Vertebrate anatomy

All

nucleus pulposus of the intervertebral discs. However, a few vertebrates, such as the sturgeon and the coelacanth, retain the notochord into adulthood.[28] 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.[29]

Fish anatomy

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, in

caudal fins, have no direct connection with the spine. They are supported by the muscles which compose the main part of the trunk.[30] 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.[31] 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 of sense organs that run along the length of the sides of fish, and these respond to nearby movements and to changes in water pressure.[30]

Sharks and rays are

oviparous and the larvae develop externally in egg cases.[32]

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.[32]

Amphibian anatomy

urinary bladder and nitrogenous waste products are excreted primarily as urea. Amphibians breathe by means of buccal pumping, 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.[33] They supplement this with gas exchange through the skin which needs to be kept moist.[34]

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.[35]

Reptile anatomy

Reptiles are a class of animals comprising turtles, tuataras, lizards, snakes and crocodiles. They are tetrapods, but the snakes and a few species of lizard 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 has evolved for internal fertilization, 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.[36]

Turtles are notable for their protective shells. They have an inflexible trunk encased in a horny

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.[37]

Tuataras superficially resemble lizards but the lineages diverged in the

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 those of other reptiles, and the lungs have a single chamber and lack bronchi. The tuatara has a well-developed parietal eye on its forehead.[37]

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.[37]

Snakes are closely related to lizards, having branched off from a common ancestral lineage during the Cretaceous period, 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.[38]

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 keratinized 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.[39]

Bird anatomy

Birds are

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.[40]

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, feet and claws on the tips of the toes.[40]

Mammal anatomy

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 specialized 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.[41]

Mammals are

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.[41]

Human anatomy

Modern anatomic technique showing sagittal sections of the head as seen by an MRI
scan
In the human, the development of skilled hand movements and increased brain size is likely to have evolved simultaneously.[42]

trunk (which includes the thorax and abdomen
), two arms and hands, and two legs and feet.

Generally, students of certain biological sciences, paramedics, prosthetists and orthotists, physiotherapists, occupational therapists, nurses, podiatrists, 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.[43]

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.[3] The major anatomy textbook, Gray's Anatomy, has been reorganized from a systems format to a regional format, in line with modern teaching methods.[44][45] A thorough working knowledge of anatomy is required by physicians, especially surgeons and doctors working in some diagnostic specialties, such as histopathology and radiology. [46]

Academic 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.[46]

Invertebrate anatomy

planktonic
crustacean

Invertebrates constitute a vast array of living organisms ranging from the simplest unicellular eukaryotes such as Paramecium to such complex multicellular animals as the octopus, lobster and dragonfly. They constitute about 95% of the animal species. By definition, none of these creatures has a backbone. The cells of single-cell protozoans have the same basic structure as those of multicellular animals but some parts are specialized 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.[47]

silica forms the exoskeleton of the microscopic diatoms and radiolaria.[48] 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.[49]

Marcello Malpighi, the father of microscopical anatomy, discovered that plants had tubules similar to those he saw in insects like the silk worm. He observed that when a ring-like portion of bark was removed on a trunk a swelling occurred in the tissues above the ring, and he unmistakably interpreted this as growth stimulated by food coming down from the leaves, and being captured above the ring.[50]

Arthropod anatomy

Arthropods comprise the largest phylum in the animal kingdom with over a million known invertebrate species.[51]

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.[53] There is considerable variation between species and many adaptations to the body parts, especially wings, legs, antennae and mouthparts.[54]

Spiders a class of arachnids have four pairs of legs; a body of two segments—a cephalothorax and an abdomen. Spiders have no wings and no antennae. They have mouthparts called chelicerae which are often connected to venom glands as most spiders are venomous. They have a second pair of appendages called pedipalps attached to the cephalothorax. These have similar segmentation to the legs and function as taste and smell organs. At the end of each male pedipalp is a spoon-shaped cymbium that acts to support the copulatory organ.

Other branches of anatomy

  • veterinary surgeons to gauge the position and anatomy of the associated deeper structures. Superficial is a directional term that indicates that structures are located relatively close to the surface of the body.[55]
  • Comparative anatomy relates to the comparison of anatomical structures (both gross and microscopic) in different animals.[3]
  • Artistic anatomy relates to anatomic studies for artistic reasons.

History

Ancient

In 1600 BCE, the

bladder, and it showed the blood vessels diverging from the heart.[56][57][58] 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.[59]

Ancient Greek anatomy and physiology underwent great changes and advances throughout the early medieval world. Over time, this medical practice expanded by a continually developing understanding of the functions of organs and structures in the body. Phenomenal anatomical observations of the human body were made, which have contributed towards the understanding of the brain, eye, liver, reproductive organs and the nervous system.

The

Hellenistic Egyptian city of Alexandria was the stepping-stone for Greek anatomy and physiology. Alexandria not only housed the biggest library for medical records and books of the liberal arts in the world during the time of the Greeks, but was also home to many medical practitioners and philosophers. Great patronage of the arts and sciences from the Ptolemy rulers helped raise Alexandria up, further rivalling the cultural and scientific achievements of other Greek states.[60]

Some of the most striking advances in early anatomy and physiology took place in Hellenistic Alexandria.

Herophilus and Erasistratus. These two physicians helped pioneer human dissection for medical research, using the cadavers of condemned criminals, which was considered taboo until the Renaissance—Herophilus was recognized as the first person to perform systematic dissections.[61] Herophilus became known for his anatomical works making impressing contributions to many branches of anatomy and many other aspects of medicine.[62] Some of the works included classifying the system of the pulse, the discovery that human arteries had thicker walls than veins, and that the atria were parts of the heart. Herophilus's knowledge of the human body has provided vital input towards understanding the brain, eye, liver, reproductive organs and nervous system, and characterizing the course of disease.[63] Erasistratus accurately described the structure of the brain, including the cavities and membranes, and made a distinction between its cerebrum and cerebellum[64] During his study in Alexandria, Erasistratus was particularly concerned with studies of the circulatory and nervous systems. He was able to distinguish the sensory and the motor nerves in the human body and believed that air entered the lungs and heart, which was then carried throughout the body. His distinction between the arteries and veins—the arteries carrying the air through the body, while the veins carried the blood from the heart was a great anatomical discovery. Erasistratus was also responsible for naming and describing the function of the epiglottis and the valves of the heart, including the tricuspid.[65] During the third century, Greek physicians were able to differentiate nerves from blood vessels and tendons[66] and to realize that the nerves convey neural impulses.[60] It was Herophilus who made the point that damage to motor nerves induced paralysis.[67] Herophilus named the meninges and ventricles in the brain, appreciated the division between cerebellum and cerebrum and recognized that the brain was the "seat of intellect" and not a "cooling chamber" as propounded by Aristotle[68] Herophilus is also credited with describing the optic, oculomotor, motor division of the trigeminal, facial, vestibulocochlear and hypoglossal nerves.[69]

Abulcasis
in the 11th century
Anatomy of the eye for the first time in history by Hunayn ibn Ishaq
in the 9th century

Great feats were made during the third century BCE in both the digestive and reproductive systems. Herophilus was able to discover and describe not only the salivary glands, but the small intestine and liver.[69] He showed that the uterus is a hollow organ and described the ovaries and uterine tubes. He recognized that spermatozoa were produced by the testes and was the first to identify the prostate gland.[69]

The anatomy of the muscles and skeleton is described in the Hippocratic Corpus, an Ancient Greek medical work written by unknown authors.[70] 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.[71][72]

In the 2nd century,

Galen of Pergamum, an anatomist, clinician, writer and philosopher,[73] wrote the final and highly influential anatomy treatise of ancient times.[74] He compiled existing knowledge and studied anatomy through dissection of animals.[73] He was one of the first experimental physiologists through his vivisection experiments on animals.[75] Galen's drawings, based mostly on dog anatomy, became effectively the only anatomical textbook for the next thousand years.[76] 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.[76]

Medieval to early modern

Anatomical study of the arm, by Leonardo da Vinci
, (about 1510)
Vesalius
's Epitome, 1543