In sciences dealing with the anatomy of animals, precise anatomical terms of location are necessary for a variety of reasons. Non-scientists often wonder why zoological and human anatomists use complex terminology to describe locations on a body, when common terms like "up", "down", "top" and "bottom" could be used.
Two major problems arise with common usage, however. First, they tend to be language-specific, requiring translation into equivalent, or almost-equivalent, terms in other languages. They are not universal terms that may be readily understood by zoologists speaking other languages. Differences in terminology remain a problem that, to some extent, still separates the fields of zoological anatomy (sometimes called zootomy) and human (medical) anatomy.
The second, and larger, problem is caused by the very nature of animals. Most animals are capable of moving relative to their environment (see Fig. 1). So while "up" might refer to the top of someone's head when they are standing upright, the same term ("up") would describe their belly while they are lying down.
Therefore, standardized anatomical (and zootomical) terms of location have been developed, usually based on Latin words, to enable all biological and medical scientists to precisely delineate and communicate information about animal (including human) bodies and their component organs.
Since animals can change position with respect to their environment, and since any appendages (arms, legs, tentacles, etc...) can change position with respect to the main body, it is important that any descriptional terms refer to the organism when it is in its standard anatomical position.
Thus, and very importantly, all descriptions are with respect to the organism in its standard anatomical position, even when the organism in question has appendages in another position. For example, see Fig. 8, where the tentacles are curved, and therefore not in anatomical position. However, a straight position is assumed when describing the proximo-distal axis. This helps avoid confusion in terminology when referring to the same organism in different postures.
There is no formal definition of standard anatomical position used in most zoology. However, the position can be loosely defined as that position in which the organism will usually be found when at rest. Thus, for most invertebrates, this would be the position in which they are normally found when not feeding, hiding, actively moving, and so on (see Figs. 6-9, below), and any appendages are straight.
For bilaterally-symmetrical organisms, such vertebrates or some invertebrates, this can be refined to include that the organisms are standing erect in a normal posture, and looking forward. (For example, see Figs. 2-4, below.)
Unlike the situation in zootomy, standard anatomical position is rigidly defined for human anatomy. As with other vertebrates, an individual is standing erect and at rest. Unlike the situation in other vertebrates, the limbs (arms and legs) are placed in unnatural positions reminiscent of the supine position adopted by cadavers during autopsy. Therefore, the individual has their feet together (or slightly separated), and their arms are rotated outward so that the palms are forward, and the thumbs are pointed away from the body. As well, the arms are usually moved slightly out from the body, so that the hands do not touch the sides.  The positions of the limbs (and the arms in particular) have important implications for directional terms in those appendages.
Ultimately, the bodies we are most familiar with are vertebrate bodies similar to or own. All vertebrates (including humans) have the same basic body plan (or bauplan) - they are bilaterally-symmetrical. That is, they have mirror-image left and right halves if divided down the centre.    For these reasons, the basic directional terms can be considered to be those used in vertebrates. By extention, the same terms are used for many other (invertebrate) organisms as well.
To begin, distinct, polar-opposite ends of ends of the organism are chosen. By definition, each pair of opposite points defines an axis. In a bilaterally-symmetrical organism, there are 6 polar opposite points, giving three axes that intersect at right angles - the x, y, and z axes familiar from three-dimensional geometry.
The most obvious end-points are the "nose" and "tail" (see Fig. 2). Anatomically, the nose is referred to as the anterior end (Latin ante; before). In organisms like vertebrates, that have distinct heads, the anterior end is sometimes be referred to as the rostral end (Latin rostrum; beak), the cranial end (Greek kranion; skull), or the cephalic end (Greek kephalē; head).   For reasons of broader applicability, especially in organisms without distinct heads (many invertebrates), anterior is usually preferred.  
The polar opposite to the anterior end is the posterior end (Latin post; after). Another term for posterior is caudal (Latin caudum; tail) — a term which strictly applies only to vertebrates, and therefore less preferred.  
By drawing a line connecting these two points, we define the anteroposterior axis (sometimes written antero-posterior). Less-used synonyms would be rostrocaudal or cephalocaudal axes (see Table 1). For brevity, the term anteroposterior is often abbreviated to read AP (or A-P) axis. As well as defining the anteroposterior axis, the terms "anterior" and "posterior" also define relative positions along the axis. Thus, in the fish in Fig. 2, the gill openings are posterior relative to the eyes, but anterior to the tail.
|Directional term||Defined Axis||Synonyms||Axis runs...|
|Anterior||Anteroposterior||Rostrocaudal1, Craniocaudal1, Cephalocaudal2||...from head end to opposite end of body or tail.|
|Dorsal||Dorsoventral||—||...from spinal column (back) to belly (front).|
|Left (lateral)||Left-right||Dextro-sinister2, Sinistro-dexter2||...from left to right sides of body.|
|Medial||Mediolateral3||—||...from centre of organism to one or other side.|
|Left or right (lateral)|
|Proximal||Proximodistal||—||...from tip of an appendage to where it joins the body.|
(1) Fairly common usage.
(2) Uncommon usage.
(3) Equivalent to one-half of the left-right axis.
(The terms "intermediate", "ipsilateral", "contralateral", "superficial" and "deep", while indicating directions, are relative terms and thus do not properly define fixed anatomical axes.)
The next most obvious end-points are the back and belly. These are termed the dorsal end (Latin dorsum; back) and the ventral end (Latin venter; abdomen), respectively. By connecting the outermostpoints the dorsoventral axis is formed (sometimes hyphenated: dorso-ventral). This is commonly abbreviated to DV (or D-V) axis. The DV axis, by definition, is perpendicular (at right angles to) the AP axis at all times (see below).
As with anteroposterior, the terms "dorsal" and "ventral" are also used to describe relative postitions along the dorsoventral axis. Thus, the pectoral fins are dorsal to the anal fin, but ventral to the dorsal fin in Fig. 2. (Note that these fins are not aligned anteroposteriorly, either - the dorsal fin being posterior to the pectoral, and anterior to the anal fins, respectively.)
The last axis, by geometric definition, must be at right angles to both the AP and DV axes. Obviously, the left side and right side of the organism are the outermost points between the two "sides" of the organism. When connected, these points form the left-right axis (commonly abbreviated to LR (or L-R) axis. Properly, this is called the dextro-sinistral (or, more uncommonly, the sinistro-dextral) axis, from the Latin dexter (right) and sinister (left). It is important to note that the "left" and "right" sides are the sides of the organism, and not that of the observer.
In practice, and contradictory to the practice with other anatomical terms of location, the vernacular "left-right" is preferentially used in English and some other languages. This is likely due to the adoption of the Latin "sinister" to mean "evil" in English and other languages (e.g. sinistre in French has the same connotation).
As with the other directions, the terms can be used as relative terms, to describe locations along the left-right axis. Thus, in Fig. 2 the dorsal fin is right of the left pectoral fin, but is left of the right eye. However, as left and right sides are mirror images, usage like this tends to be somewhat confusing, as structures are duplicated on both sides (i.e. above there is both a right eye and a left eye, forcing one to specify which is used as a reference).
To counter this clumsiness of usage, the directional term lateral (Latin lateralis; "to the side") is used as a modifier for both sides, yielding the left lateral and right lateral sides. As an opposite to lateral, the term medial (Latin medius; "middle") is used to define a point in the centre of the organism (where the left-right axis intersects the midsagittal plane — see below). Thus, rather than "left-right" axis and its inherent clumsiness of usage, the term mediolateral (also sometimes hyphenated medio-lateral) axis is frequently used. Sometimes this is abbreviated to ML (or M-L) axis.   Properly, the ML axis is a half axis; practically, its usage is less clumsy and less linguistically biased than "left-right". The terms may still be used relatively to describe locations along the LR axis. Thus, in Fig. 2 the gills are medial to the operculum, but lateral to the heart.
The usage "mediolateral" is strictly used to describe relative position along the left-right axis, to avoid confusion with the terms "superficial" and "deep" (see below).
Together, the AP, DV and LR (or ML) axes allow for precise three-dimensional descriptions of location within any bilaterally-symmetrical organism, whether vertebrate or invertebrate. In practice, the terms can cause some confusion when, unlike the fish shown in Fig. 2, the organism in question is not strictly linear in form (see Figs. 3 and 4). For example, the AP axis in Fig. 3 does not appear to be at right angles to the DV axis. Rather, it is a depiction of the approximate average AP axis, when all body segments are included.
When considering any one segment, the dorsoventral axis is perpendicular to the AP axis. Thus, in Fig. 4, the DV axis of the tail would run from the "back" of the tail (posterior end of the trunk), to the "underside" of the tail (near the legs) — nearly parallel to the AP axis of the main body.
As a general rule of thumb, if the body is included in consideration, the AP axis of the main body would be used, as would the DV and ML axes perpendicular to it. However, if considering only one segment, the AP axis would shift to reflect the axes shown in Fig. 4, with the DV and ML axes shifting correspondingly. Alternatively, to avoid confusion, AP, DV and ML terms are used strictly in relation to the main body, and the terms proximal and distal are used for body segments such as the head, neck and tail (see below).
As noted above, the standard AP, DV and ML directional axes, can cause some confusion when describing parts of the body that can change position (move) relative to the main body. This is particularly true when considering appendages. "Appendages" would include vertebrate fins (see Fig. 2) and limbs (see Figs. 3 and 4), but properly apply to any structure that extends (and can at least potentially move separately) from the main body. Thus, "appendage" would also include such structures as external ears (pinneae) and hair (in mammals), feathers (in birds) and scales (fish, reptiles and birds). As well, varieties of tentacles or and other projections from the body in invertebrates and the male penis in many vertebrates and some invertebrates, would be included.
Since appendages often move independently of (and therefore change position with respect to) the main body, separate directional terms are used when describing them. The term proximal (Latin proximus; nearest) is used to describe where the appendage joins the body, and the term distal (Latin distare; to stand away from) is used for the point furthest from the point of attachment to the body.
By connecting the two points, the proximodistal (sometimes hyphenated to proximo-distal) axis. (The abbreviation AB axis is occasionally, but not commonly, used.) As before, the terms "proximal" and "distal" can be used as relative terms to indicate where structures lie along the proximodistal axis. Thus, the "elbow" is proximal to the hoof, but distal to the "shoulder" in Figs. 3 and 4.
Chosing terms for the other two axes perpendicular to the proximodistal axis could be variable, as they would also depend on the position of the limb. For that reason, when considering any organism, the other two axes are considered to be relative to the appendage when in standard anatomical position. This is roughly defined for all organisms, as in the normal position when at rest and not moving. For tetrapod vertebrates, this includes the caveat that they are standing erect and not lying down. Thus, the fish in Fig. 2, and the horse in Figs. 3 and 4 are in standard anatomical position. (Special considerations with respect to limb position are applied in human anatomy — see below).
In addition to the three primary axes (AP, DV and the ML half-axis) and the proximodistal axis of appendages, several directional terms can be used in bilaterally symmetrical animals. Some of these terms were These terms are strictly relative, and as such do not and cannot be used to define fixed axes. These terms include:
The large variety of body shapes present in invertebrates presents a difficult problem when attempting to apply standard directional terms. As such, some terms are taken directly from the vertebrate terms by analogy, and appropriate novel terms are applied, as necessary. In all cases, the usage of terms is dependent on the bauplan of the organism.
The large variety of body shapes present in invertebrates presents a difficult problem when attempting to apply standard directional terms. Depending on the organism, some terms are taken by analogy from the vertebrate terms, and appropriate novel terms are applied, as necessary. In all cases, the usage of terms is dependent on the bauplan of the organism.
In organisms with a changeable shape, such as amoeboid organisms (Fig. 5a), directional terms are meaningless, since the shape of the organism is changeable, and no fixed axes are present. Similarly, in organisms that are spherical in shape (Fig. 5b), an infinite number of axes, all passing through the centre are possible. Each axis is functionally and practically indistinguishable from all others, and therefore meaningless. In such organisms, only the terms superficial and deep hold any descriptive meaning.
In organisms that maintain a constant shape and have one dimension longer than the other, at least two directional terms can be used. The long or longitudinal axis is defined by points at the opposite ends of the organism. Similarly, a perpendicular transverse axis is defined by points on the opposite sides of the organism. No third axis can usually be defined in such organisms. Usually such organisms, like that pictured in Fig. 6, are microscopic, protists, aquatic, and planktonic (not attached to a surface).
Some elongated protists have distinctive ends of the body. In such organisms, the end with a mouth (or equivalent structure, such as the cytostome in Paramecium or Stentor), or the end that that usually points in the direction the organism moves (such as the end with the flagella in Euglena can be designated as the anterior end. The opposite end then becomes the posterior end, and by connecting them, an anteroposterior axis is formed. Properly, this terminology would only apply to an organism that is always planktonic (not normally attached to a surface — as in Fig. 6 top), although the term can also be applied to one that is sessile (normally attached to a surface — as in Fig. 6, bottom and Fig. 7).
In elongated organisms, the distinctive shorter axis is usually referred to as the transverse axis. Again, no third axis is usually possible, although there are exceptions, particularly where a non-terminal cytostome or other unique structure is present.
Organisms that are sttached to a substrate, such as sponges (Fig. 7), or some animal-like protists also have distinctive ends. The part of the organism attached to a substrate is usually referred to as the basal end (Latin basis; support or foundation), whereas the end furthest from the attachment is referred to as the apical end (Latin apex; peak, tip). Thus, by joining the two ends, an apical-basal (or basal-apical) axis is formed (see Fig. 7). Again, the shorter dimension would be the transverse axes, and multiple equivalent axes are present as there is no symmetry present.
Radially symmetrical organisms include those in the group Radiata — primarily jellyfish, sea anemones and corals and the comb jellies.  Adult echinoderms (sea stars (starfish), sea urchins, and sea cucumbers and others) are also included, since they are pentaradial (i.e. they have 5 radial axes of symmetry). Echinoderm larvae are not included, since they are bilaterally-symmetrical. 
Unlike spherical and asymmetrical organisms, radially-symmetrical animals always have one distinctive axis.
Cnidarians have an incomplete digestive system, meaning that one end of the organism has a mouth, and the opposite end has no opening from the gut (coelenteron). For this reason, the end of the organism with the mouth is referred to as the oral end (Latin oris; mouth), and the opposite surface is the aboral end (Latin ab-; prefix meaning "away from"). Thus, by joining the polar opposite oral and aboral ends, an oral-aboral axis is formed (Fig. 8).
|Bilateral Bauplans||Radial Bauplans|
|Anterior||Rostral, Cranial, Cephalic2||Oral||Apical3|
(1) Includes both Radiates and adult Echinoderms.
(2) Rarely used.
(3) Only in organisms attached to a substrate.
(4) Vertebrate equivalents are meaningless in radial animals.
(5) Roughly equivalent to "superficial".
(6) Roughly equivalent to "deep".
As with vertebrates, appendages that move independently of the body (tentacles in cnidarians and comb jellies), have a definite proximodistal axis (Fig. 8). Unlike vertebrates, cnidarians (jellyfish, sea anemones, corals) have no other distinctive axes, and multiple radial axes are possible (Fig. 9).
It is noteworthy that some "biradially-symmetrical" comb jellies have distinct "tentacular" and "pharyngeal" axes,  and are thus anatomically equivalent to bilaterally-symmetrical animals.) As well, adult echinoderms (starfish, sea urchins, sea cucumbers) are pentaradial, and have only five symmetrical radial axes (unlike the multiple axes in cnidarians).
"Lateral", dorsal, and ventral have no meaning in such organisms, and all can be replaced by the generic term peripheral (Latin peri-; around; see Table 2). Medial can be used, but in the case of radiates indicates the central point of these organisms, rather than a central axis (as in vertebrates). Thus, as there are many possible radial axes, there are multiple medio-peripheral (half-) axes (Fig. 9).
As bilaterally-symmetrical organisms, anatomical directions in humans can correctly be described using the same terms as those for vertebrates and other members of the taxonomic group Bilateria. However, for historical and other reasons, standard human directional terminology has several differences from that used for other bilaterally-symmetrical organisms.
Although it can be argued that the standard directional nomenclature used for vertebrate zootomy can and should be used for medical anatomy, the differences persist. The differences in terminology arose (and are perpetuated) three primary reasons:
Unfortunately, the persistence of medical terminology as distinct from that used for other vertebrates tends to be confusing. For a quick comparison of equivalent terminology used in vertebrate and human anatomy, see Table 3 (below).
|Vertebrate zootomy||Human anatomy|
|Anterior||Rostral, Cranial, Cephalic1||Superior||Same1|
(1) Rarely used.
(2) Used only to describe one side of an appendage.
(3) Strictly relative term, used with other locational descriptors.
As with other vertebrates, two of the most obvious extremes are the "top" and the "bottom" of the organism. In standard anatomical position, these equate to the head and feet, respectively in humans. The head end is referred to as the superior end (Latin superior: "above"), while the feet are referred to as the inferior end (Latin inferior: "below"). Thus, the axis formed by joining the two is the superior-inferior axis. .
As in other vertebrates, there are synonymous terms for superior and inferior (Table 3). The terms cranial, cephalic, and rostral are occasionally encountered. "Cranial", as a reference to the skull, is fairly commonly used, whereas "cephalic" is uncommonly used, and "rostral" is extremely rarely used in human antaomy. Similarly, the term caudal is occasionally used in human anatomy, and the cranio-caudal axis is occasionally encountered. Generally, this usage would only be used with respect to the head and main body (trunk), and not when considering the limbs.
Anterior and posterior, as used in medical/human anatomical descriptions are major sources of confusion to those accustomed to standard vertebrate directional terminology, and vice versa. The confusion arises from the differences in standard anatomical positions of quadruped verebrates and bipedal humans.
In human anatomical usage, anterior refers to the "front" of the individual, and is synonymous with ventral. Similarly, posterior, in medical anatomy refers to the "back" of the subject, and is synonymous with dorsal (see Table 3). The terms "dorsal" and "ventral" are used in human anatomy, but infrequently when referring to the body as a whole. Thus, the anteroposterior axis is preferred usage for describing the axis connecting the front and the back in humans. 
Left and right lateral are used in the same sense as they are in other vertebrates, as is medial. The left-right axis is rarely used in medicine, however, the mediolateral axis is used almost exclusively. 
As in other vertebrates, the terms "proximal" and "distal" are used to describe the point of attachment to, and part of an appendage furthest away from, the body, respectively. However, other terms are used for direction in the appendages, given the unique position of the limbs (in standard anatomical position) in humans.
In standard anatomical position, the palms of the hands point anteriorly. Thus, anterior can be (and sometimes is) used to the palm of the hand, and posterior can be (and sometimes is) used to describe the back of the hand and arm.
However, presumably for improved clarity, the directional term palmar (Latin palma; palm of the hand) is usually used for the anterior of the hand, and dorsal is used to describe the back of the hand. Thus, by connecting the extremes, dorsopalmaraxis is formed. Most commonly, "dorsopalmar" is used when describing the hand, although it is sometimes applied to the arm as a whole (see Fig. 11).
For the third axis, the mediolateral axis suffices, although if referring to the limb alone, "medial" may refer to the centre of the arm itself.
Specialized terms are used to describe location on appendages, parts that have a point of attachment to the main trunk of the body. Structures that are close to the point of attachment of the body are proximal or central, while ones more distant from the attachment point are distal or peripheral. For example, the hands are at the distal end of the arms, while the shoulders are at the proximal ends. These terms can also be used relatively to organs, for example the proximal end of the urethra is attached to the bladder.
In the limbs of most animals, the terms cranial and caudal are used in the regions proximal to the carpus (the wrist, in the forelimb) and the tarsus (the ankle in the hindlimb). Objects and surfaces closer to or facing towards the head are cranial; those facing away or further from the head are caudal.
Nearer the carpal joint, the term dorsal replaces cranial and palmar replaces caudal. Similarly, nearer the tarsal joint the term dorsal replaces cranial and plantar replaces caudal. For example, the top of a dog's paw is its dorsal surface; the underside, either the palmar (on the forelimb) or the plantar (on the hindlimb) surface.
The sides of the forearm are named after its bones: Structures closer to the radius are radial, structures closer to the ulna are ulnar, and structures relating to both bones are referred to as radioulnar. Similarly, in the lower leg, structures near the tibia (shinbone) are tibial and structures near the fibula are fibular (or peroneal).
Volar (sometimes used as a synonym for "palmar") refers to the underside, for both the palm and the sole (plantar), as in volar pads on the underside of hands, fingers, feet and toes.
The terms valgus and varus are used to refer to angulation of the distal part of a limb at a joint. For example, at the elbow joint, in the anatomical position, the forearm and the upper arm do not lie in a straight line, but the forearm is angulated laterally with respect to the upper arm by about 5-10°. The forearm is said to be "in valgus". Angulation at a joint may be normal (as in the elbow) or abnormal.
Three basic reference planes are used in zoological anatomy.
For post-embryo humans a coronal plane is vertical and a transverse plane is horizontal, but for embryos and quadripeds a coronal plane is horizontal and a transverse plane is vertical.
When describing anatomical motion, these planes describe the axis along which an action is performed. So by moving through the transverse plane, movement travels from head to toe. For example, if a person jumped directly up and then down, their body would be moving in the transverse plane.
Some of these terms come from Latin. Sagittal means "like an arrow," a reference to the position of the spine which naturally divides the body into right and left equal halves, the exact meaning of the term "midsagittal".
Sometimes the orientation of certain planes needs to be distinguished, for instance in medical imaging techniques such as sonography, CT scans, MRI scans or PET scans. One imagines a human in the anatomical position, and an X-Y-Z coordinate system with the X-axis going from front to back, the Y-axis going from left to right, and the Z-axis going from up to down. The X-axis axis is always forward (Tait-Bryan angles) and the right-hand rule applies.
The axes and the sagittal plane are the same for bipeds and quadrupeds, but the orientation of the coronal and transverse planes switch. The axes on particular pieces of equipment may or may not correspond to axes of the body, especially since the body and the equipment may be in different relative orientations.
In discussing the neuroanatomy of animals, particularly rodents used in neuroscience research, the convention has been to name the sections of the brain according to the homologous human sections. Hence, what is technically a transverse section with respect the body of a rat (dividing anterior from posterior) may often be referred to in rat neuroanatomical coordinates as a coronal section, and likewise a coronal section with respect to the body (ie. dividing ventral from dorsal) in a rat brain is referred to as transverse. This preserves the comparison with the human brain which is rotated with respect to the body axis by 90 degrees in the ventral direction. It does mean that the planes of the rat brain are not necessarily the same as those of the body.
In humans, reference may be made to landmarks which are on the skin or visible underneath. As with planes, lines and points are imaginary. Examples include:
Additionally, reference may be made to structures at specific levels of the spine (e.g. the 4th cervical vertebra, abbreviated "C4"), or the rib cage (e.g. the 5th intercostal space, abbreviated "5ICS").
Anatomical terms are applied to the subject (human or animal) in the standard anatomical position. This is typically a (standing) posture similar to that seen during life, rather than lying on a table. For humans, the body is standing erect, feet together and toes pointed forward, arms at the sides and palms facing forward (forearms supine). In other species, e.g. quadrupeds, the standard anatomical position is described as standing erect with the head facing forwards in a neutral position.
In humans, the anatomical position of the skull has been agreed by international convention to be the Frankfurt plane, a position where the lower margins of the orbits and the upper margins of the ear canals all lie in the same horizontal plane. This is a good approximation to the position where the subject is standing upright and facing forwards.
The content of this section is licensed under the GNU Free Documentation License (local copy). It uses material from the Wikipedia article "Anatomical terms of location " modified December 8, 2007 with previous authors listed in its history.