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For the anatomy of plants, seePlant anatomy.For other uses, seeAnatomy (disambiguation).

One of the large, detailed illustrations inAndreas Vesalius'sDe humani corporis fabrica16th century, marking the rebirth of anatomy

Anatomy(Greekanatomē,'dissection') is the branch ofbiologyconcerned with the study of the structure oforganismsand their parts.[1]Anatomy is a branch of natural science which deals with the structural organization of living things. It is an old science, having its beginnings in prehistoric times.[2]Anatomy is inherently tied todevelopmental biology,embryology,comparative anatomy,evolutionary biology,andphylogeny,[3]as these are the processes by which anatomy is generated, both over immediate and long-term timescales. Anatomy andphysiology,which study the structure andfunctionof organisms and their parts respectively, make a natural pair ofrelated disciplines,and are often studied together.Human anatomyis one of the essentialbasic sciencesthat areappliedinmedicine.[4]

The discipline of anatomy is divided intomacroscopicandmicroscopic.Macroscopic anatomy, orgross anatomy,is the examination of an animal's body parts using unaidedeyesight.Gross anatomy also includes the branch ofsuperficial anatomy.Microscopic anatomy involves the use of optical instruments in the study of thetissuesof various structures, known ashistology,and also in the study ofcells.

Thehistory of anatomyis characterized by a progressive understanding of the functions of theorgansand structures of the human body. Methods have also improved dramatically, advancing from the examination of animals bydissectionof carcasses andcadavers(corpses) to 20th centurymedical imagingtechniques includingX-ray,ultrasound,andmagnetic resonance imaging.

Definition

A dissected body, lying prone on a table, byCharles Landseer

Derived from theGreekἀνατομήanatomē"dissection" (fromἀνατέμνωanatémnō"I cut up, cut open" from ἀνάaná"up", and τέμνωtémnō"I cut" ),[5]anatomy is the scientific study of the structure oforganismsincluding their systems, organs andtissues.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 fromphysiologyandbiochemistry,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 ofbile,the role of the liver in nutrition and the regulation of bodily functions.[6]

The discipline of anatomy can be subdivided into a number of branches including gross ormacroscopicanatomy andmicroscopicanatomy.[7]Gross anatomyis the study of structures large enough to be seen with the naked eye, and also includessuperficial anatomyor surface anatomy, the study by sight of the external body features.Microscopic anatomyis the study of structures on a microscopic scale, along withhistology(the study of tissues), andembryology(the study of an organism in its immature condition).[3]

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 includedissection,in which a body is opened and its organs studied, andendoscopy,in which avideo camera-equipped instrument is inserted through a small incision in the body wall and used to explore the internal organs and other structures.AngiographyusingX-raysormagnetic resonance angiographyare methods to visualize blood vessels.[8][9][10][11]

The term "anatomy" is commonly taken to refer tohuman anatomy.However, substantially the same structures and tissues are found throughout the rest of the animal kingdom and the term also includes the anatomy of other animals. The termzootomyis 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 inplant anatomy.[6]

Animal tissues

Stylized cutaway diagram of an animal cell (with flagella)

ThekingdomAnimaliacontainsmulticellular organismsthat areheterotrophicandmotile(although some have secondarily adopted asessilelifestyle). Most animals have bodies differentiated into separatetissuesand these animals are also known aseumetazoans.They have an internaldigestivechamber, with one or two openings; thegametesare produced in multicellular sex organs, and thezygotesinclude ablastulastage in theirembryonic development.Metazoans do not include thesponges,which have undifferentiated cells.[12]

Unlikeplant cells,animal cellshave neither a cell wall norchloroplasts.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 cell, including those found inmuscles,nervesandskin.Each typically has a cell membrane formed ofphospholipids,cytoplasmand anucleus.All of the different cells of an animal are derived from the embryonicgerm layers.Those simpler invertebrates which are formed from two germ layers of ectoderm and endoderm are calleddiploblasticand the more developed animals whose structures and organs are formed from three germ layers are calledtriploblastic.[13]All of a triploblastic animal's tissues and organs are derived from the three germ layers of the embryo, theectoderm,mesodermandendoderm.

Animal tissues can be grouped into four basic types:connective,epithelial,muscleandnervous tissue.

Hyaline cartilage at high magnification (H&E stain)

Connective tissue

Connective tissuesare fibrous and made up of cells scattered among inorganic material called theextracellular matrix.Connective tissue gives shape to organs and holds them in place. The main types are loose connective tissue,adipose tissue,fibrous connective tissue,cartilageandbone.The extracellular matrix containsproteins,the chief and most abundant of which iscollagen.Collagen plays a major part in organizing and maintaining tissues. The matrix can be modified to form askeletonto support or protect the body. Anexoskeletonis a thickened, rigidcuticlewhich is stiffened bymineralization,as incrustaceansor by the cross-linking of its proteins as ininsects.Anendoskeletonis internal and present in all developed animals, as well as in many of those less developed.[13]

Epithelium

Gastric mucosa at low magnification (H&E stain)

Epithelial tissueis composed of closely packed cells, bound to each other bycell adhesion molecules,with little intercellular space. Epithelial cells can besquamous(flat),cuboidalorcolumnarand rest on abasal lamina,the upper layer of thebasement membrane,[14]the lower layer is the reticular lamina lying next to the connective tissue in the extracellular matrix secreted by the epithelial cells.[15]There are many different types of epithelium, modified to suit a particular function. In therespiratory tractthere is a type ofciliatedepithelial lining; in the small intestine there aremicrovillion the epithelial lining and in the large intestine there areintestinal villi.Skinconsists of an outer layer ofkeratinizedstratified squamous epithelium that covers the exterior of the vertebrate body.Keratinocytesmake up to 95% of the cells in theskin.[16]The epithelial cells on the external surface of the body typically secrete an extracellular matrix in the form of acuticle.In simple animals this may just be a coat ofglycoproteins.[13]In more advanced animals, manyglandsare formed of epithelial cells.[17]

Muscle tissue

Cross section throughskeletal muscleand a smallnerveat high magnification (H&E stain)

Muscle cells(myocytes) form the active contractile tissue of the body.Muscle tissuefunctions to produce force and cause motion, either locomotion or movement within internal organs. Muscle is formed of contractilefilamentsand is separated into three main types;smooth muscle,skeletal muscleandcardiac muscle.Smooth muscle has nostriationswhen examined microscopically. It contracts slowly but maintains contractibility over a wide range of stretch lengths. It is found in such organs assea anemonetentacles and the body wall ofsea cucumbers.Skeletal 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 inearthwormsthat can extend slowly or make rapid contractions.[18]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 walls of theuterus,bladder,intestines,stomach,oesophagus,respiratory airways,andblood vessels.Cardiac muscleis found only in theheart,allowing it to contract and pump blood round the body.

Nervous tissue

See also:Neuroanatomy

Nervous tissueis composed of many nerve cells known asneuronswhich transmit information. In some slow-movingradially symmetricalmarine animals such asctenophoresandcnidarians(includingsea anemonesandjellyfish), the nerves form anerve net,but in most 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, specialized receptor cells such aschemoreceptorsandphotoreceptorsare found in groups and send messages alongneural networksto other parts of the organism. Neurons can be connected together inganglia.[19]In higher animals, specialized receptors are the basis of sense organs and there is acentral nervous system(brain and spinal cord) and aperipheral nervous system.The latter consists ofsensory nervesthat transmit information from sense organs andmotor nervesthat influence target organs.[20][21]The peripheral nervous system is divided into thesomatic nervous systemwhich conveys sensation and controlsvoluntary muscle,and theautonomic nervous systemwhich involuntarily controlssmooth muscle,certain glands and internal organs, including thestomach.[22]

Vertebrate anatomy

See also:Comparative anatomy
Mouseskull

Allvertebrateshave a similar basicbody planand at some point in their lives, mostly in theembryonicstage, share the majorchordatecharacteristics; a stiffening rod, thenotochord;a dorsal hollow tube of nervous material, theneural tube;pharyngeal arches;and a tail posterior to the anus. Thespinal cordis protected by thevertebral columnand is above the notochord and thegastrointestinal tractis below it.[23]Nervous tissue is derived from theectoderm,connective tissues are derived frommesoderm,and gut is derived from theendoderm.At the posterior end is atailwhich continues the spinal cord and vertebrae but not the gut. The mouth is found at the anterior end of the animal, and theanusat the base of the tail.[24]The defining characteristic of a vertebrate is thevertebral column,formed in the development of the segmented series ofvertebrae.In most vertebrates the notochord becomes thenucleus pulposusof theintervertebral discs.However, a few vertebrates, such as thesturgeonand thecoelacanthretain the notochord into adulthood.[25]Jawed vertebratesare typified by paired appendages, fins or legs, which may be secondarily lost. The limbs of vertebrates are considered to behomologousbecause the same underlying skeletal structure was inherited from their last common ancestor. This is one of the arguments put forward byCharles Darwinto support his theory ofevolution.[26]

Fish anatomy

Main article:Fish anatomy
Cutaway diagram showing various organs of a fish

The body of afishis 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, incartilaginous fish,or bone inbony fish.The main skeletal element is the vertebral column, composed of articulatingvertebraewhich are lightweight yet strong. The ribs attach to the spine and there are nolimbsor limb girdles. The main external features of the fish, thefins,are composed of either bony or soft spines called rays, which with the exception of thecaudal fins,have no direct connection with the spine. They are supported by the muscles which compose the main part of the trunk.[27]The heart has two chambers and pumps the blood through the respiratory surfaces of thegillsand on round the body in a single circulatory loop.[28]The eyes are adapted for seeing underwater and have only local vision. There is an inner ear but no external ormiddle ear.Low frequency vibrations are detected by thelateral linesystem 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.[27]

Sharks and rays arebasalfish with numerousprimitiveanatomical 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 dermalplacoid scales.They have acloacainto which the urinary and genital passages open, but not aswim bladder.Cartilaginous fish produce a small number of large,yolkyeggs. Some species areovoviviparousand the young develop internally but others areoviparousand the larvae develop externally in egg cases.[29]

The bony fish lineage shows morederivedanatomical 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 anoperculum,and a mouth at or near the tip of the snout. The dermis is covered with overlappingscales.Bony fish have a swim bladder which helps them maintain a constant depth in the water column, but not a cloaca. They mostlyspawna large number of small eggs with little yolk which they broadcast into the water column.[29]

Amphibian anatomy

Main article:Amphibian anatomy
Frog skeleton
Skeleton ofSurinam horned frog(Ceratophrys cornuta)
Plastic model of a frog

Amphibiansare aclassof animals comprisingfrogs,salamandersandcaecilians.They aretetrapods,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 havearticular 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 littlekeratinand lacks scales, but contains manymucous glandsand in some species, poison glands. The hearts of amphibians have three chambers, twoatriaand oneventricle.They have aurinary bladderandnitrogenous waste productsare excreted primarily asurea.Amphibians breathe by means ofbuccal pumping,a pump action in which air is first drawn into thebuccopharyngealregion through the nostrils. These are then closed and the air is forced into the lungs by contraction of the throat.[30]They supplement this withgas exchangethrough the skin which needs to be kept moist.[31]

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 resembleearthwormsand 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.[32]

Reptile anatomy

Main article:Reptile anatomy
Skeleton of adiamondback rattlesnake

Reptilesare a class of animals comprisingturtles,tuataras,lizards,snakesandcrocodiles.They aretetrapods,but the snakes and a few species oflizardeither 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 theirlungsby 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 acopulatory organpresent in most species. The eggs are surrounded byamniotic membraneswhich prevents them from drying out and are laid on land, ordevelop internallyin some species. The bladder is small as nitrogenous waste is excreted asuric acid.[33]

Turtlesare notable for their protective shells. They have an inflexible trunk encased in a hornycarapaceabove and aplastronbelow. 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.[34]

Tuatarassuperficially resemble lizards but the lineages diverged in theTriassicperiod. 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 those of other reptiles, and the lungs have a single chamber and lackbronchi.The tuatara has a well-developedparietal eyeon its forehead.[34]

Lizardshave skulls with only onefenestraon 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.[34]

Snakesare closely related to lizards, having branched off from a common ancestral lineage during theCretaceousperiod, 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 ofpelvic 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.[35]

Crocodiliansare 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.[36]

Bird anatomy

Main article:Bird anatomy
Part of a wing.Albrecht Dürer,c. 1500–1512

Birdsaretetrapodsbut though their hind limbs are used for walking or hopping, their front limbs arewingscovered withfeathersand adapted for flight. Birds areendothermic,have a highmetabolic rate,a lightskeletal systemand powerfulmuscles.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.[37]

The feathers are outgrowths of theepidermisand 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 singleuropygial glandnear the base of the tail. This produces an oily secretion that waterproofs the feathers when the birdpreens.There are scales on the legs, feet and claws on the tips of the toes.[37]

Mammal anatomy

Main article:Mammal anatomy

Mammalsare 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 ofprismatic enamel.The teeth are shed once (milk teeth) during the animal's lifetime or not at all, as is the case incetaceans.Mammals have three bones in the middle ear and acochleain theinner ear.They are clothed in hair and their skin contains glands which secretesweat.Some of these glands are specialized asmammary glands,producing milk to feed the young. Mammals breathe withlungsand have a musculardiaphragmseparating 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.[38]

Mammals areamniotes,and most areviviparous,giving birth to live young. The exception to this are the egg-layingmonotremes,theplatypusand theechidnasof Australia. Most other mammals have aplacentathrough which the developingfoetusobtains nourishment, but inmarsupials,the foetal stage is very short and the immature young is born and finds its way to its mother'spouchwhere it latches on to anippleand completes its development.[38]

Human anatomy

Further information:Human body § Human anatomy,andOutline of human anatomy
Modern anatomic technique showing sagittal sections of the head as seen by aMRIscan
In the human, the development of skilled hand movements and increased brain size is likely to have evolved simultaneously.[39]

Humans have the overall body plan of a mammal. Humans have ahead,neck,trunk(which includes thethoraxandabdomen), twoarmsandhands,and twolegsandfeet.

Generally, students of certainbiological sciences,paramedics,prosthetists and orthotists,physiotherapists,occupational therapists,nurses,podiatrists,andmedical studentslearn 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 ofdissectionand inspection ofcadavers.The study of microscopic anatomy (orhistology) can be aided by practical experience examining histological preparations (or slides) under amicroscope. [40]

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.[41][42]A thorough working knowledge of anatomy is required by physicians, especiallysurgeonsand doctors working in some diagnostic specialties, such ashistopathologyandradiology. [43]

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.[43]

Invertebrate anatomy

Head of a maleDaphnia,aplanktoniccrustacean

Invertebratesconstitute a vast array of living organisms ranging from the simplest unicellulareukaryotessuch asParameciumto such complex multicellular animals as theoctopus,lobsteranddragonfly.They constitute about 95% of the animal species. By definition, none of these creatures has a backbone. The cells of single-cellprotozoanshave 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 byciliaorflagellaor may proceed via the advance ofpseudopodia,food may be gathered byphagocytosis,energy needs may be supplied byphotosynthesisand the cell may be supported by anendoskeletonor anexoskeleton.Some protozoans can form multicellular colonies.[44]

Metazoansare a multicellular organism, with different groups of cells serving different 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 anextracellular matrixwhich provides support to the organism. An endoskeleton derived from themesodermis present inechinoderms,spongesand somecephalopods.Exoskeletonsare derived from the epidermis and is composed ofchitininarthropods(insects, spiders, ticks, shrimps, crabs, lobsters).Calcium carbonateconstitutes the shells ofmolluscs,brachiopodsand some tube-buildingpolychaete wormsandsilicaforms the exoskeleton of the microscopicdiatomsandradiolaria.[45]Other invertebrates may have no rigid structures but the epidermis may secrete a variety of surface coatings such as thepinacodermof sponges, the gelatinous cuticle of cnidarians (polyps,sea anemones,jellyfish) and thecollagenouscuticle ofannelids.The outer epithelial layer may include cells of several types including sensory cells, gland cells and stinging cells. There may also be protrusions such asmicrovilli,cilia, bristles,spinesandtubercles.[46]

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.[47]

Arthropod anatomy

Main articles:Arthropod,Insect morphology,andSpider anatomy

Arthropodscomprise the largest phylum in the animal kingdom with over a million known invertebrate species.[48]

Insectspossesssegmentedbodies supported by a hard-jointed outer covering, theexoskeleton,made mostly of chitin. The segments of the body are organized into three distinct parts, a head, athoraxand anabdomen.[49]The head typically bears a pair of sensoryantennae,a pair ofcompound eyes,one to three simple eyes (ocelli) and three sets of modified appendages that form themouthparts.The thorax has three pairs of segmentedlegs,one pair each for the three segments that compose the thorax and one or two pairs ofwings.The abdomen is composed of eleven segments, some of which may be fused and houses thedigestive,respiratory,excretoryand reproductive systems.[50]There is considerable variation between species and many adaptations to the body parts, especially wings, legs, antennae and mouthparts.[51]

Spidersa class ofarachnidshave four pairs of legs; a body of two segments—acephalothoraxand anabdomen.Spiders have no wings and no antennae. They have mouthparts calledcheliceraewhich are often connected to venom glands as most spiders are venomous. They have a second pair of appendages calledpedipalpsattached 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 thecopulatory organ.

Other branches of anatomy

  • Superficial or surface anatomyis important as the study of anatomical landmarks that can be readily seen from the exterior contours of the body.[3]It enables physicians orveterinary surgeonsto 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.[52]
  • Comparative anatomyrelates to the comparison of anatomical structures (both gross and microscopic) in different animals.[3]
  • Artistic anatomy relates to anatomic studies for artistic reasons.

History

Main article:History of anatomy

Ancient

Image of early rendition of anatomy findings

In 1600 BCE, theEdwin Smith Papyrus,anAncient Egyptianmedical text,described theheart,its vessels,liver,spleen,kidneys,hypothalamus,uterusandbladder,and showed theblood vesselsdiverging from the heart. TheEbers 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.[53]

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.

TheHellenistic Egyptiancity ofAlexandriawas 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 thePtolemyrulers helped raise Alexandria up, further rivalling the cultural and scientific achievements of other Greek states.[54]

An anatomythangka,part ofDesi Sangye Gyatso's The Blue Beryl, 17th century

Some of the most striking advances in early anatomy and physiology took place in Hellenistic Alexandria.[54]Two of the most famous anatomists and physiologists of the third century wereHerophilusandErasistratus.These two physicians helped pioneer humandissectionfor medical research. They also conductedvivisectionson the cadavers of condemned criminals, which was considered taboo until the Renaissance—Herophilus was recognized as the first person to perform systematic dissections.[55]Herophilus became known for his anatomical works making impressing contributions to many branches of anatomy and many other aspects of medicine.[56]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.[57]Erasistratus accurately described the structure of the brain, including the cavities and membranes, and made a distinction between its cerebrum and cerebellum[58]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.[59]During the third century, Greek physicians were able to differentiate nerves from blood vessels and tendons[60]and to realize that the nerves convey neural impulses.[54]It was Herophilus who made the point that damage to motor nerves induced paralysis.[61]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[62]Herophilus is also credited with describing the optic, oculomotor, motor division of the trigeminal, facial, vestibulocochlear and hypoglossal nerves.[63]

Surgical instruments were invented for the first time in history byAbulcasisin the 11th century
Anatomy of the eye for the first time in history byHunayn ibn Ishaqin the 9th century
13th century anatomical illustration

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.[63]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.[63]

The anatomy of the muscles and skeleton is described in theHippocratic Corpus,an Ancient Greek medical work written by unknown authors.[64]Aristotledescribedvertebrateanatomy based on animaldissection.Praxagorasidentified the difference betweenarteriesandveins.Also in the 4th century BCE,HerophilosandErasistratusproduced more accurate anatomical descriptions based onvivisectionof criminals inAlexandriaduring thePtolemaic dynasty.[65][66]

In the 2nd century,Galen of Pergamum,ananatomist,clinician,writer andphilosopher,[67]wrote the final and highly influential anatomy treatise of ancient times.[68]He compiled existing knowledge and studied anatomy through dissection of animals.[67]He was one of the first experimental physiologists through hisvivisectionexperiments on animals.[69]Galen's drawings, based mostly on dog anatomy, became effectively the only anatomical textbook for the next thousand years.[70]His work was known toRenaissancedoctors only throughIslamic Golden Agemedicine until it was translated from the Greek some time in the 15th century.[70]

Medieval to early modern

Anatomical study of the arm, byLeonardo da Vinci,(about 1510)
Anatomical chart inVesalius'sEpitome,1543
Michiel Jansz van Mierevelt–Anatomy lesson of Dr. Willem van der Meer,1617

Anatomy developed little from classical times until the sixteenth century; as the historian Marie Boas writes, "Progress in anatomy before the sixteenth century is as mysteriously slow as its development after 1500 is startlingly rapid".[70]: 120–121 Between 1275 and 1326, the anatomistsMondino de Luzzi,Alessandro AchilliniandAntonio BenivieniatBolognacarried out the first systematic human dissections since ancient times.[71][72][73]Mondino'sAnatomyof 1316 was the first textbook in the medieval 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.[70]

Leonardo da Vinci(1452–1519) was trained in anatomy byAndrea del Verrocchio.[70]He made use of his anatomical knowledge in his artwork, making many sketches of skeletal structures, muscles and organs of humans and other vertebrates that he dissected.[70][74]

Andreas Vesalius(1514–1564) (Latinized from Andries van Wezel), professor of anatomy at theUniversity of Padua,is considered the founder of modern human anatomy.[75]Originally fromBrabant,Vesalius published the influential bookDe humani corporis fabrica( "the structure of the human body" ), a large format book in seven volumes, in 1543.[76]The accurate and intricately detailed illustrations, often inallegoricalposes against Italianate landscapes, are thought to have been made by the artistJan van Calcar,a pupil ofTitian.[77]

In England, anatomy was the subject of the first public lectures given in any science; these were given by theCompany of Barbers and Surgeonsin the 16th century, joined in 1583 by the Lumleian lectures in surgery at theRoyal College of Physicians.[78]

Late modern

Further information:History of anatomy in the 19th century

In the United States, medical schools began to be set up towards the end of the 18th century. Classes in anatomy needed a continual stream of cadavers for dissection and these were difficult to obtain. Philadelphia, Baltimore and New York were all renowned forbody snatchingactivity as criminals raided graveyards at night, removing newly buried corpses from their coffins.[79]A similar problem existed in Britain where demand for bodies became so great that grave-raiding and evenanatomy murderwere practised to obtain cadavers.[80]Some graveyards were in consequence protected with watchtowers. The practice was halted in Britain by theAnatomy Actof 1832,[81][82]while in the United States, similar legislation was enacted after the physicianWilliam S. ForbesofJefferson Medical Collegewas found guilty in 1882 of "complicity with resurrectionists in the despoliation of graves in Lebanon Cemetery".[83]

The teaching of anatomy in Britain was transformed by SirJohn Struthers,Regius Professor of Anatomyat theUniversity of Aberdeenfrom 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 ofcomparative anatomy,published over 70 research papers, and became famous for his public dissection of theTay Whale.[84][85]From 1822 the Royal College of Surgeons regulated the teaching of anatomy in medical schools.[86]Medical museums provided examples in comparative anatomy, and were often used in teaching.[87]Ignaz Semmelweisinvestigatedpuerperal feverand he discovered how it was caused. He noticed that the frequently fatal fever occurred more often in mothers examined by medical students than by midwives. The students went from the dissecting room to the hospital ward and examined women in childbirth. Semmelweis showed that when the trainees washed their hands in chlorinated lime before each clinical examination, the incidence of puerperal fever among the mothers could be reduced dramatically.[88]

An electron microscope from 1973

Before the modern medical era, the main means for studying the internal structures of the body weredissectionof the dead andinspection,palpationandauscultationof the living. It was the advent ofmicroscopythat opened up an understanding of the building blocks that constituted living tissues. Technical advances in the development ofachromatic lensesincreased theresolving powerof the microscope and around 1839,Matthias Jakob SchleidenandTheodor Schwannidentified that cells were the fundamental unit of organization of all living things. Study of small structures involved passing light through them and themicrotomewas invented to provide sufficiently thin slices of tissue to examine. Staining techniques using artificial dyes were established to help distinguish between different types of tissue. Advances in the fields ofhistologyandcytologybegan in the late 19th century[89]along with advances in surgical techniques allowing for the painless and safe removal ofbiopsyspecimens. The invention of theelectron microscopebrought a great advance in resolution power and allowed research into theultrastructureof cells and theorganellesand other structures within them. About the same time, in the 1950s, the use ofX-ray diffractionfor studying the crystal structures of proteins, nucleic acids and other biological molecules gave rise to a new field ofmolecular anatomy.[89]

Equally important advances have occurred innon-invasivetechniques for examining the interior structures of the body.X-rayscan be passed through the body and used in medicalradiographyandfluoroscopyto differentiate interior structures that have varying degrees of opaqueness.Magnetic resonance imaging,computed tomography,andultrasound imaginghave all enabled examination of internal structures in unprecedented detail to a degree far beyond the imagination of earlier generations.[90]

See also

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Bibliography

Library resourcesabout
Anatomy
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