Anatomical terms of location

Standard anatomical terms of location are employed in science which deal with the anatomy of animals to avoid ambiguities which might otherwise arise. They are not language-specific, and thus require no translation. They are universal terms that may be readily understood by zoologists who speak any language.

While these terms are standardized within specific fields of biology, they can differ dramatically from one discipline to another. Differences in terminology remain a problem that, to some extent, still separates the fields of zoological anatomy (sometimes called zootomy) and human (medical) anatomy (sometimes called androtomy).

The Craniata (vertebrates) share a substantial heritage of common structure, allowing much of the same terminology to be used for all of them. It is necessary for this terminology to be based on the anatomy of the animal in a standard way to avoid ambiguities such as might occur if a word such as "top" were used, which might designate the head of a human but the left or right side of a flounder. Most animals, furthermore, are capable of moving relative to their environment. So while "up" might refer to the direction of a standing human's head, the same term ("up") might be used to refer to the direction of the belly of a supine human. It is also necessary to employ some specific anatomical knowledge in order to apply the terminology unambiguously: E.g. while the ears would be superior to (above) the shoulders in a human, this fails when describing the armadillo, where the shoulders are above the ears. Thus in veterinary terminology, the ears would be cranial to (i.e. "towards the head from") the shoulders in the armadillo, the dog, the kangaroo, or any other vertebrate, including the human. Similarly, while the belly is considered anterior to (in front of) the back in humans, this terminology fails for the flounder, the armadillo and the dog. In veterinary terms, the belly would be ventral ("towards the abdomen") in all vertebrates. In human anatomy, as will be explained below, all naming is based on positions relative to the body in a standing (standard anatomical) position with arms at the side and palms facing forwards (thumbs out). While the universal vertebrate terminology used in veterinary medicine would work in human medicine, the human terms are thought to be too well established to change.

For invertebrates, locational terminology becomes more complicated, as many species are not bilaterally symmetrical. In these species, terminology depends on the type of symmetry present (if any).

Thus, 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.

Standard anatomical position

Because animals can change orientation with respect to their environment, and because appendages (arms, legs, tentacles, etc.) can change position with respect to the main body, it is important that positional descriptive terms refer to the organism when it is in its standard anatomical position.

Thus, 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. 9, 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.

Invertebrate and vertebrate zootomy

Medical (human) anatomy

Unlike the situation in zootomy, standard anatomical position is rigidly defined for human anatomy. As with other vertebrates, the human body is standing erect and at rest. Unlike the situation in other vertebrates, the limbs are placed in positions reminiscent of the supine position imposed on cadavers during autopsy. Therefore, the body has its feet together (or slightly separated), and its arms are rotated outward so that the palms are forward, and the thumbs are pointed away from the body (forearms supine). As well, the arms are usually moved slightly out from the body, so that the hands do not touch the sides.^[1][2] The positions of the limbs (and the arms in particular) have important implications for directional terms in those appendages. The penis in males is also erect in the anatomical position, hence the dorsal surface of the penis is actually anterior in the flaccid state.^[3]

Skull

In humans, the anatomical position of the skull has been agreed by international convention to be the Frankfurt plane, a position in which the lower margins of the orbits, the orbitales, and the upper margins of the ear canals, the poria, all lie in the same horizontal plane. This is a good approximation to the position in which the skull would be if the subject were standing upright and facing forward normally.

Directional terms

Ultimately, the bodies we are most familiar with are vertebrate bodies similar to our 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.^[4][5][6][7] For these reasons, the basic directional terms can be considered to be those used in vertebrates. By extension, the same terms are used for many other (invertebrate) organisms as well.

Vertebrate directional terms

To begin with, distinct, polar-opposite 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.

Figure 2: Anatomical directions and defined axes in a vertebrate

Anterior and posterior

"Anterior" and "Posterior" redirect here. For other uses, see Anterior (disambiguation) and Posterior (disambiguation).

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 referred to as the rostral end (Latin rostrum; beak), the cranial end (Greek kranion; skull), or the cephalic end (Greek kephalē; head).^[4][5][7] For reasons of broader applicability, especially in organisms without distinct heads (many invertebrates), "anterior" is usually preferred.^[5][8][9]

The polar opposite to the anterior end is the posterior end (Latin post; after). Another term for posterior is caudal (Latin caudum; tail, though in humans this refers to the feet i.e. inferior rather than posterior)—a term which strictly applies only to vertebrates, and therefore less preferred, except in veterinary medicine where these terms are standard.^{[5][8][9][10]}

By drawing a line connecting these two points, we define the anteroposterior axis (sometimes written antero-posterior). Caudal and Posterior (back end) are often used interchangeably. In veterinary medicine, caudo-cranial is preferred between head and tail, and rostro-caudal between nose and neck. 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.

Table 1: Defined Axes in Vertebrate Zoology

Directional term	Defined Axis	Synonyms	Axis runs...
Anterior	Anteroposterior	Rostrocaudal1, Craniocaudal1, Cephalocaudal2	...from head end to opposite end of body or tail.
Posterior
Dorsal	Dorsoventral	—	...from spinal column (back) to belly (front).
Ventral
Left (lateral)	Left-right	Dextro-sinister2, Sinistro-dexter2	...from left to right sides of body.
Right (lateral)
Medial	Mediolateral3	—	...from centre of organism to one or other side.
Left or right (lateral)
Proximal	Proximodistal	—	...from tip of an appendage (distal) to where it joins the body (proximal).
Distal
Notes: (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. Also, while the "rostrocaudal" and anteroposterior directionality are equivalent in a significant portion of the human body, they are different directions in other parts of the body.)

Dorsal and ventral

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 outermost points 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 positions 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.)

Left and right (lateral), and medial

The last axis, by geometric definition, must be at right angles to both the AP and DV axes. 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. In Latin, this is called the dextro-sinistral (or, more uncommonly, the sinistro-dextral) axis, from dexter (right) and sinister (left). It is important to note that the "left" and "right" sides are the sides of the organism, and not those of the observer.

"Left-right" is typically used in English and some other languages.^{[clarification needed]}

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 median (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), and the term medial means "towards the median plane". 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.^[4][5][7] 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).

Sources of confusion

Figure 3: Directional axes in the tetrapod vertebrate Equus caballus (a horse). The axis between cranial and caudal is the Cr-Cd axis, and between the dorsal and ventral is the D-V axis. (Left-right axis not shown; image shows the right side of the organism.)

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, which includes most tetrapods (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.

Figure 4: Different directional AP axes in three body segments of a horse). Axis (A) (in red) shows the AP axis of the tail, (B) shows the AP axis of the neck, and (C) shows the AP axis of the head.

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).

To avoid this confusion, in veterinary medicine, the terms anterior, posterior, superior, and inferior are generally avoided except for certain structures within the head.^[10] By using the terms cranial, caudal, dorsal and ventral, all tetrapod organisms (including bipeds) can be described uniformly.

Rostral and Caudal shown on a human skull

In humans, the directions "rostral" and "caudal" often become confused with anterior and posterior, or superior and inferior. The difference between the two is most easily visualized when looking at the head, as can be seen in the image to the right. From the most caudal of positions in the nervous system (of a person) to a nearby, rostral area, it is equally accurate to say the area in question is rostral as to say it is superior. However, in the frontal lobes of the telencephalon, to say an area is rostral to a nearby area is equivalent to saying it is anterior (or ventral). Those two lines lie on planes perpendicular to one another. This occurs, as becomes clear in the diagram, due to the intuitive yet curious curving "C" shape of rostrocaudal directionality when discussing the human brain.

Proximal and distal

The term proximal (Latin proximus; nearest) describes 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. Since appendages often move independently of (and therefore change position with respect to) the main body, these separate directional terms are used when describing them.

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 (pinnae) and hair (in mammals), feathers (in birds) and scales (fish, reptiles and birds). As well, varieties of tentacles or other projections from the body in invertebrates and the male in many vertebrates and some invertebrates, would be included.

By connecting the two points, the proximodistal (sometimes hyphenated to proximo-distal) axis is created. (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.

Choosing 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).

Ostial and distal

Similar to appendages which branch out from the body, the directions of blood vessels may be labeled with the terms Ostial (referring to the Ostium or opening where the vessel branches off) and distal (the extreme end away from the branch point).

Teeth

Teeth may be aligned with their main axis identical to that of the jaw, but they can also be rotated. In such a case using terms like "anterior" or "lateral" can be confusing. Therefore, a special set of terms exists, mainly used in palaeontology.^[11]

• lingual ([Latin] lingua; tongue): on the side that faces the tongue.
• labial ([Latin] labium; lip): facing the lips
• buccal ([Latin] bucca; cheek): on the side that faces the cheek.

• mesial: toward the mandibular symphysis (where lower jaws meet)
• distal: away from the mandibular symphysis

Dentistry uses the same terminology, as well as

• vestibular ([Portugese] vestibulo; entrance): the side away from the tongue (includes labial and buccal).

Other directional terms

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. These terms are strictly relative, and as such do not and cannot be used to define fixed axes. These terms include:

• Ipsilateral (Latin ipse; self/same + latus-eris, "side"): on the same side as another structure. Thus, the left arm is ipsilateral to the left leg.
• Contralateral (Latin contra; against): on the opposite side to another structure. Thus, the left arm is contralateral to the right arm, or the right leg.
• Superficial (Latin superfacies; at the surface or face): near the outer surface of the organism. Thus, skin is superficial to the muscle layer. The opposite is "deep", or "visceral".
• Deep: further away from the surface of the organism. Thus, the muscular layer is deep to the skin, but superficial to the intestines. This is one of the few terms where the English vernacular is prevalent. The proper anglicised Latin term would be profound (Latin profundus; due to depth), but this word has other meanings in English. In other languages, the equivalent term is usually similar to "profound" (e.g. profond, meaning deep, in French).
• Intermediate (Latin intermedius; inter, between and medius, middle): between two other structures. Thus, the navel is intermediate to (or intermediate between) the left arm and the contralateral (right) leg.
• Visceral (Latin viscus,-eris; internal organs, flesh): associated with organs within the body's cavities. The stomach is a viscus within the abdominal cavity, and is covered with a lining called the visceral peritoneum.
• Parietal (Latin paries,-etis, "wall"): pertaining to the wall of a body cavity. The parietal peritoneum is the lining on the inside of the abdominal cavity. (Parietal can also refer specifically to the parietal bone of the skull or associated structures.)
• Axial (Latin axis from Greek axōn "axle"): Towards the central axis of the organism or the extremity.
• Abaxial : away from the central axis of the organism or extremity
• Rostral (Latin - rostr(um), beak or nose ): situated toward the oral or nasal region, or in the case of the brain, towards the tip of the frontal lobe.
• Caudal (Latin - cauda, tail): of, at, or near the tail or the posterior end of the body. In the human case, towards the bottom of the feet (also the "tail" of the spinal cord, and body).

Invertebrate directional terms

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.

Figure 5: Asymmetrical and spherical body shapes. (a) An organism with an asymmetrical bauplan (Amoeba proteus—an amoeba). (b) An organism with a spherical bauplan (Actinophrys sol—a heliozoan.

Asymmetrical and spherical organisms

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), there is nothing to distinguish one line through the centre of the organism from another. An infinite number of triads of mutually perpendicular axes could be defined, but any such choice of axes would be functionally and practically indistinguishable from all others, and therefore would be useless. In such organisms, only the terms superficial and deep hold any descriptive meaning.

Figure 6: Four individuals of Phaeodactylum tricornutum, a diatom with a fixed elongated shape.

Elongated organisms

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 can be defined by points on opposite sides of the organism. There is typically no basis for the definition of a third axis. Usually such organisms, like that pictured in Fig. 6, are planktonic (free-swimming) protists, and are nearly always viewed on microscope slides, where they appear essentially two-dimensional. In some cases a third axis can be defined, particularly where a non-terminal cytostome or other unique structure is present.^[9]

Elongated organisms with distinctive ends

Figure 7: Organisms where the ends of the long axis are distinct. (Paramecium caudatum, above, and Stentor roeseli, below.)

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 usually points in the direction of the organism's locomotion (such as the end with the flagellum in Euglena), is normally designated as the anterior end. The opposite end then becomes the posterior end, and by connecting them, an anteroposterior axis is formed.^[9] Properly, this terminology would only apply to an organism that is always planktonic (not normally attached to a surface, as in Fig. 7 top), although the term can also be applied to one that is sessile (normally attached to a surface, as in Fig. 7, bottom, and Fig. 8).^[12]

Figure 8: A cluster of Euplectella aspergillum sponges (Venus flower baskets), showing the apical-basal axes.

Organisms that are attached to a substrate, such as sponges (Fig. 8), or some animal-like protists also have distinctive ends. The part of the organism attached to the 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. 8). Transverse axes may be defined indifferently in any direction perpendicular to this axis, as there is no symmetry present.

Radially-symmetrical organisms

Radially symmetrical organisms include those in the group Radiata—primarily jellyfish, sea anemones and corals and the comb jellies.^[5][9] Adult echinoderms (sea stars (starfish), sea urchins, and sea cucumbers and others) are also included, since they are pentaradial (i.e. they have fivefold discrete rotational symmetry). Echinoderm larvae are not included, since they are bilaterally symmetrical.^[5][9]

Figure 9: Chrysoara spp. (a jellyfish), showing the oral-aboral, and proximodistal axes. (Note that the appendages are not in standard anatomical position, so that the axis is curved.)

Unlike spherical and asymmetrical organisms, radially-symmetrical animals always have one distinctive axis.

Figure 10: Aurelia aurita, another species of jellyfish, showing multiple radial and medio-peripheral axes.

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).^[9] 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. 9).

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. 10).

It is noteworthy that some "biradially-symmetrical" comb jellies have distinct "tentacular" and "pharyngeal" axes,^[13] 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. 10).

Table 2: Comparison of Directional Terms used in Radially-Symmetrical1 and Bilaterally-Symmetrical Animals
Bilateral Bauplans		Radial Bauplans
Direction	Synonyms	Direction	Synonyms
Anterior	Rostral, Cranial, Cephalic2	Oral	Apical3
Posterior	Caudal2	Aboral	Basal3
Dorsal	—	Peripheral4,5	—
Ventral	—	Peripheral4,5	—
Left (lateral)	Sinister	Peripheral4,5	—
Right (lateral)	Dexter	Peripheral4,5	—
Medial	—	Same6	—
Proximal	—	Same	—
Distal	—	Same	—
Notes: (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".

Arachnids

Two specialized terms are sometimes used for describing views of arachnid legs and pedipalps. Prolateral refers to the surface of a leg which is closest to the anterior end of an arachnid's body. Retrolateral refers to the surface of a leg which is closest to the posterior end of an arachnid's body.^[14]

Medical (human) directional terms

As humans are bilaterally-symmetrical organisms, anatomical directions in humans can usually be correctly 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.

Why zootomy and androtomy terms differ

The terms of zootomy and androtomy came into usage at a time when all scientific communication took place in Latin. In their original Latin forms the respective meanings of "anterior" and "posterior" are in front of (or before) and behind (or after), those of "dorsal" and "ventral" are towards the spine and towards the belly, and those of "superior" and "inferior" are above and below. From these meanings it can be seen that in the most general terms the anterior/posterior axis is oriented to the direction of forward motion, the dorsal/ventral axis is oriented to the anatomy of the vertebrate torso, and the superior/inferior axis is oriented to gravity.

For almost all vertebrates, including almost all bipeds, these axes all provide a consistent reference for anatomical positions across species—with the inferior/superior axis being roughly the same as the dorsal/ventral axis, and therefore redundant. Humans, however, have the rare property of having a torso oriented perpendicular to their direction of forward motion—while their head orientation remains consistent with other vertebrates on this axis. This makes the dorsal/ventral axis on humans redundant with the anterior/posterior axis, and the inferior/superior axis necessary. Because of this difference with humans, the anterior/posterior and inferior/superior axes are inconsistent between humans and other vertebrates in torso anatomy but consistent in head anatomy. As all three of these axes are used in the naming of anatomical structures, and most human anatomical structures are shared by other animals, these differences can lead to considerable confusion. For example, in the naming of brain structures, the non-human context of the dorsal/ventral axis was used. Therefore, in human anatomy, "dorsal" can refer to two different (perpendicular) directions—the posterior direction in the context of the torso, and the superior direction in the context of the brain. Ironically, the "dorsal" direction in the human brain, besides being perpendicular to the "dorsal" direction in the human torso, is actually the opposite direction of what might be inferred from the literal Latin meaning of "toward the spine".^{[citation needed]}

While it would be possible to introduce a system of axes that is completely consistent between humans and other vertebrates by having two separate pairs of axes, one used exclusively for the head (e.g. anterior/posterior and inferior/superior) and the other exclusively for the torso (e.g. dorsal/ventral and caudal("toward the tail")/rostral("toward the beak")), doing so would require the renaming of very many anatomical structures.

For a quick comparison of equivalent terminology used in vertebrate and human anatomy, see Table 3 (below).

Table 3: Equivalent directional terms used in
vertebrate zoology and human anatomy

Vertebrate zootomy		Human torso		Human head
Direction	Synonyms	Direction	Synonyms	Direction	Synonyms
Anterior	Rostral, Cranial, Cephalic1	Superior	Same1, Up	Anterior	Front
Posterior	Caudal	Inferior	Caudal1, Down	Posterior	Back
Dorsal	—	Posterior	Dorsal, Back	Superior	Dorsal, Up
Ventral	—	Anterior	Ventral, Front	Inferior	Ventral, Down
lateral	Away from the middle	Same	—	Same	—
Left (lateral)	Sinister1	Same	—	Same	—
Right (lateral)	Dexter1	Same	—	Same	—
Medial	Middle	Same	—	Same	—
Proximal	Away from extremity	Same	—	Same	—
Distal	Towards extremity	Same	—	Same	—
Intermediate2	—	Same	—	Same	—
Ipsilateral2	Same side	Same	—	Same	—
Contralateral2	Opposite side	Same	—	Same	—
Superficial2	—	Same	—	Same	—
Deep2	—	Same	—	Same	—
Notes: (1) Rarely used. (2) Strictly relative term, used with other locational descriptors.

Superior and inferior

As with other vertebrates, two of the most obvious extremes are the "top" and the "bottom" of the organism. In standard anatomical position, these correspond 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.^[15][16]

As with other vertebrate terminology, there are synonymous terms for superior and inferior (Table 3). The terms cranial and cephalic are often encountered. "Cranial", as a reference to the skull, is fairly commonly used, whereas "cephalic" is uncommonly used. The term "rostral" is rarely used in human anatomy, referring more to the front of the face than the superior aspect of the organism. This term is more applicable in organisms with longer heads, such as equids.^[17] Similarly, the term caudal is occasionally used in human anatomy,^[17] 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.

As with vertebrate directional terms, superior and inferior can be used in a relative sense in humans, but can not be uniformly applied to other organisms with varying normal anatomical positions. For example, the shoulders are superior to the navel, but inferior to the eyes in humans. In any tetrapod, the shoulders are cranial to the belly, but caudal to the eyes.

Anterior and posterior

In human anatomical usage, anterior refers to the "front" of the individual, and is synonymous with ventral, other than in the head. Similarly, posterior, refers to the "back" of the subject, and is synonymous with dorsal, other than in the head (see Table 3).^[18] The terms "dorsal" and "ventral" are used in human anatomy, but infrequently when referring to the body as a whole.^[19] The anteroposterior axis is preferred usage for describing the axis connecting the front and the back in humans.^[18][20]

"Anterior" and "posterior" can also be used as relative terms. Thus, the eyes are posterior to the nose, but anterior to the back of the head in humans.
However, in the horse, for example, the eyes are caudal to the nose, and rostral to the back of the head.

Left and right (lateral), and medial

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; instead, the mediolateral axis is used almost exclusively.^[17][21]

Appendages

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.

Arms

Figure 12: The directional terms used in a human hand.

In standard anatomical position, the palms of the hands point anteriorly. Thus, anterior can be used to describe the palm of the hand, and posterior can be 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, dorsopalmar axis is formed. Most commonly, "dorsopalmar" is used when describing the hand, although it is sometimes applied to the arm as a whole (see Fig. 12).

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.

Relative directions

Also, in common usage, the segments of the digestive system closest to the mouth are termed proximal, as opposed to those closest to the anus, which are termed distal. The terms oral "of the mouth" and aboral "away from the mouth" are also used.

Relative directions in the limbs

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 farther from the head are caudal.

Distal to the carpal joint, the term dorsal replaces cranial and palmar replaces caudal. Similarly, distal to 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.

Planes

Anatomical planes in a human

General usage

Three basic reference planes are used in zoological anatomy.

• A sagittal plane, being a plane parallel to the sagittal suture, divides the body into sinister and dexter (left and right) portions.
  • The midsagittal or median plane is in the mid line; i.e. it would pass through mid line structures such as the navel or spine, and all other sagittal planes (also referred to as parasagittal planes) are parallel to it. Median can also refer to the midsagittal plane of other structures, such as a digit.
• A coronal or frontal plane divides the body into dorsal and ventral (back and front, or posterior and anterior) portions.
• A transverse plane, also known as an axial plane or cross-section, divides the body into cranial and caudal (head and tail) portions.

For post-embryonic humans a coronal plane is vertical and a transverse plane is horizontal, but for embryos and quadrupeds 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 through the transverse plane in the coronal and sagittal planes.

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", or to the shape of the sagittal suture, which defines the sagittal plane and is shaped like an arrow.

A longitudinal plane is any plane perpendicular to the transverse plane. The coronal plane and the sagittal plane are examples of longitudinal planes.

Usage in human anatomy

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 Z-axis going from front to back, the X-axis going from left to right, and the Y-axis going from up to down. The Z-axis axis is always forward (Tait-Bryan angles) and the right-hand rule applies.

• A transverse (also known as axial or horizontal) plane is an X-Z plane, parallel to the ground, which (in humans) separates the superior from the inferior, or put another way, the head from the feet.
• A coronal (also known as frontal) plane is a Y-X plane, perpendicular to the ground, which (in humans) separates the anterior from the posterior, the front from the back, the ventral from the dorsal.
• A sagittal (also known as lateral) plane is an Y-Z plane, perpendicular to the ground, which separates left from right. The midsagittal plane is the specific sagittal plane that is exactly in the middle of the body.

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.

Occasionally, in medicine, abdominal organs may be described with reference to the trans-pyloric plane which is a transverse plane passing through the pylorus.

Anatomical planes in animal brains

In discussing the neuroanatomy of animals, particularly rodents used in neuroscience research, a simplistic convention has been to name the sections of the brain according to the homologous human sections. Hence, what is technically a transverse (orthogonal) section with respect to the body length axis 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 (i.e. dividing ventral from dorsal) in a rat brain is referred to as transverse. This preserves the comparison with the human brain, whose length axis in rough approximation is rotated with respect to the body axis by 90 degrees in the ventral direction. It implies that the planes of the brain are not necessarily the same as those of the body.

Actually, the situation is more complex, since comparative embryology shows that the length axis of the neural tube (the primordium of the brain) has three internal bending points, namely two ventral bendings at the cervical and cephalic flexures (cervical flexure roughly between the medulla oblongata and the spinal cord, and cephalic flexure between the diencephalon and the midbrain), and a dorsal (pontine or rhombic) flexure at the midst of the hindbrain, behind the cerebellum. The latter flexure mainly appears in mammals and sauropsids (reptiles and birds), whereas the other two, and principally the cephalic flexure, appear in all vertebrates (the sum of the cervical and cephalic ventral flexures is the cause of the 90 degree angle mentioned above in humans between body axis and brain axis). This more realistic concept of the longitudinal structure of vertebrate brains implies that any section plane, except the sagittal plane, will intersect variably different parts of the same brain as the section series proceeds across it (relativity of actual sections with regard to topological morphological status in the ideal unbent neural tube). Any precise description of a brain section plane therefore has to make reference to the anteroposterior part of the brain to which the description refers (e.g., transverse to the midbrain, or horizontal to the diencephalon). A necessary note of caution is that modern embryologic orthodoxy indicates that the brain's true length axis finishes rostrally somewhere in the hypothalamus where basal and alar zones interconnect from left to right across the median line; therefore, the axis does not enter the telencephalic area, although various authors, both recent and classic, have assumed a telencephalic end of the axis. The causal argument for this lies in the end of the axial mesoderm -mainly the notochord, but also the prechordal plate- under the hypothalamus. Early inductive effects of the axial mesoderm upon the overlying neural ectoderm is the mechanism that establishes the length dimension upon the brain primordium, jointly with establishing what is ventral in the brain (close to the axial mesoderm) in contrast with what is dorsal (distant from the axial mesoderm). Apart of the lack of a causal argument for introducing the axis in the telencephalon, there is the obvious difficulty that there is a pair of telencephalic vesicles, so that a bifid axis is actually implied in these outdated versions.

Surface and other landmarks in humans

Axillary lines.

In humans, reference may take origin from superficial anatomy, made to landmarks which are on the skin or visible underneath. As with planes, lines and points are imaginary. Examples include:

• The midaxillary line, a line running vertically down the surface of the body passing through the apex of the axilla (armpit). Parallel are the anterior axillary line, which passes through the anterior axillary skinfold, and the posterior axillary line, which passes through the posterior axillary skinfold.
• The mid-clavicular line, a line running vertically down the surface of the body passing through the midpoint of the clavicle.
• The mid-pupillary line, a line running vertically down the face through the midpoint of the pupil when looking directly forwards.
• The mid-inguinal point, a point midway between the anterior superior iliac spine and the pubic symphysis.
  • mid-point of inguinal ligament = mid-point between anterior superior iliac spine and pubic tubercle
• Tuffier's line, which is a transverse line passing across the lumbar spine between the posterior iliac crests.
• Mid-ventral line, the intersection between the ventral skin and the median plane.

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).

See also

• Anatomical terms of motion
• Commonly used terms of relationship and comparison in dentistry
• Human positions

Notes

1. ^ Marieb (1995), pp. 13–14.
2. ^ Tortora and Derrickson (2006), pp. 12–13
3. ^ Faculty of Biological Science, University of Leeds: Introductory Anatomy http://www.leeds.ac.uk/chb/lectures/anatomy2.html
4. ^ ^{a b c} Kardong (2005).
5. ^ ^{a b c d e f g} Hickman et al. (2003).
6. ^ Houseman (2003).
7. ^ ^{a b c} Wischnitzer (1993).
8. ^ ^{a b} Miller (2002).
9. ^ ^{a b c d e f g} Ruppert et al. (2004).
10. ^ ^{a b} Nomina Anatomica Veterinaria
11. ^ Smith, J.B. and Dodson, P. (2003) "A proposal for a standard terminology of anatomical notation and orientation in fossil vertebrate dentitions." Journal of Vertebrate Paleontology 23:1-12.
12. ^ Valentine, James W. (2004). On the Origin of Phyla. Chicago: University of Chicago Press. ISBN 0226845486. 
13. ^ Ruppert et al. (2004), p. 184.
14. ^ Kaston, B. J. (1972). How to Know the Spiders (2nd ed.). Dubuque, Iowa: W. C. Brown Co. p. 19. ISBN 0697048993. 
15. ^ Marieb (1995)
16. ^ Tortora and Derrickson (2006)
17. ^ ^{a b c} Tortora and Derrickson (2006), p. 14.
18. ^ ^{a b} Tortora and Derrickson (2006) p. 14.
19. ^ The term "dorsal" is used with respect to limb position, however.
20. ^ Marieb (1995) p. 16
21. ^ Marieb (1995), p. 16.

References

• Atkins, B.T., Duval, A., Lewis, H.M.A., and Milne, R.C. (1993) Collins-Robert French-English, English-French Dictionary, 2nd ed. HarperCollins, Glasgow UK and Le Robert, Paris, France. ISBN 0-06-275513-7.
• Barber, K., ed. (1998) The Canadian Oxford English Dictionary. Oxford University Press, Don Mills, ON, Canada. ISBN 0-19-541120-X.
• Campbell, N.A. and Reece, J.B. (2005) Biology, 7th ed. Pearson, San Francisco, CA, USA. ISBN 0-8053-7166-4.
• Hickman, C.P., Jr., Roberts, L.S. and Larson, A. (2003) Animal Diversity, 3rd ed. McGraw-Hill, New York, NY, USA. ISBN 0-07-234903-4.
• Houseman, J. (2003) Digital Zoology, v. 2.0. McGraw-Hill, New York, NY, USA. ISBN 0-07-256481-4.
• Kardong, K. (2005) Vertebrates: Comparative Anatomy, Function, Evolution, 4th ed. McGraw-Hill, New York, NY, USA. ISBN 0-07-290956-0.
• Marieb, E.N. (1995) Human Anatomy and Physiology, 3rd ed. Benjamin/Cummings, Redwood City, CA, USA. ISBN 0-8053-4281-8.
• Miller, S.A. (2002) General Zoology Laboratory Manual, 5th ed. McGraw-Hill, New York, NY, USA. ISBN 0-07-250837-0 (Mix from ISBN 0-07-243559-3).
• Ruppert, E.E., Fox, R.S. and Barnes, R.D. (2004) Invertebrate Zoology: A Functional Evolutionary Approach, 7th ed. Thomson, Belmont, CA, USA. ISBN 0-03-025982-7.
• Tortora, G.J. and Derrickson, B. (2006) Principles of Anatomy and Physiology, 11th ed. Wiley, Hoboken, NJ, USA. ISBN 0-471-68934-3.
• Wischnitzer, S. (1993) Atlas and Dissection Guide for Comparative Anatomy, 5th ed. W.H. Freeman, New York. ISBN 0-71-672374-3.

External links

• Anatomy of the lower limb
• Directions, cross sections, and divisions of the human brain