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In
anatomy Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...
a chiasm is the spot where two structures cross, forming an X-shape (). Examples of chiasms are: * A tendinous chiasm, the spot where two
tendons A tendon or sinew is a tough band of dense fibrous connective tissue that connects muscle to bone. It sends the mechanical forces of muscle contraction to the skeletal system, while withstanding tension. Tendons, like ligaments, are made of ...
cross. For example, the tendon of the
flexor digitorum superficialis muscle Flexor digitorum superficialis (''flexor digitorum sublimis'') or flexor digitorum communis sublimis is an extrinsic flexor muscle of the fingers at the proximal interphalangeal joints. It is in the anterior compartment of the forearm. It is s ...
, and the tendon of the
flexor digitorum longus muscle The flexor digitorum longus muscle or flexor digitorum communis longus is situated on the tibial side of the leg. At its origin it is thin and pointed, but it gradually increases in size as it descends. It serves to flex the second, third, fourth ...
which even forms two chiasms. * In
neuroanatomy Neuroanatomy is the study of the structure and organization of the nervous system. In contrast to animals with radial symmetry, whose nervous system consists of a distributed network of cells, animals with bilateral symmetry have segregated, defi ...
, the crossing of fibres of a
nerve A nerve is an enclosed, cable-like bundle of nerve fibers (called axons). Nerves have historically been considered the basic units of the peripheral nervous system. A nerve provides a common pathway for the Electrochemistry, electrochemical nerv ...
or the crossing of two nerves.


Neural chiasms

Different types of crossings of nerves are referred to as chiasm: * Type I: Two nerves can ''cross one over the other'' (
sagittal plane The sagittal plane (; also known as the longitudinal plane) is an anatomical plane that divides the body into right and left sections. It is perpendicular to the transverse and coronal planes. The plane may be in the center of the body and divi ...
) without fusing, e.g., the
trochlear nerve The trochlear nerve (), ( lit. ''pulley-like'' nerve) also known as the fourth cranial nerve, cranial nerve IV, or CN IV, is a cranial nerve that innervates a single muscle - the superior oblique muscle of the eye (which operates through the pu ...
(see figure). * Type II: Two nerves can ''merge'' while at least part of the fibres cross the midline (see figure 2). * Type III: The fibres ''within'' a single nerve cross, such that the order of the functional map is reversed, e.g., the optic chiasms of various
invertebrates Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...
such as
insects Insects (from Latin ') are hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of jointed ...
and
cephalopods A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
. * Type IV: A torsion or loop by 180 degrees of a nerve can also reverse the order of the functional map. This type is usually not referred to as chiasm. Note that in the third type there is no crossing of the mid sagittal plane. Only in the first type, the crossing is complete. There are other kinds of crossings of nerve fibres. The chiasm is distinguished from a
decussation Decussation is used in biological contexts to describe a crossing (due to the shape of the Roman numeral for ten, an uppercase 'X' (), ). In Latin anatomical terms, the form is used, e.g. . Similarly, the anatomical term Chiasm (anatomy), chi ...
, which is a crossing of nerve fibres inside the
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain, spinal cord and retina. The CNS is so named because the brain integrates the received information and coordinates and influences the activity o ...
. A chiasm also differs from a
ganglion A ganglion (: ganglia) is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system, this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system, there are ...
in that axons run through it without making any synapses. A chiasm is thus not a nervous processing centre.


Optic chiasm in vertebrates

By far the most widely known chiasm is the optic chiasm in vertebrate animals, including humans.


Examples

Chiasms are found in vertebrates but also in invertebrates. The optic chiasm in vertebrates can be of type I or II. However, an optic chiasm of type III is found in many
insects Insects (from Latin ') are hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of jointed ...
and in
cephalopods A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
. In
vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
, three of the
cranial nerves Cranial nerves are the nerves that emerge directly from the brain (including the brainstem), of which there are conventionally considered twelve pairs. Cranial nerves relay information between the brain and parts of the body, primarily to and f ...
show a chiasm. *The optic chiasm of the
optic tract In neuroanatomy, the optic tract () is a part of the visual system in the brain. It is a continuation of the optic nerve that relays information from the optic chiasm to the ipsilateral lateral geniculate nucleus (LGN), pretectal nuclei, and su ...
(type I or II) *The chiasm of the
Trochlear nerve The trochlear nerve (), ( lit. ''pulley-like'' nerve) also known as the fourth cranial nerve, cranial nerve IV, or CN IV, is a cranial nerve that innervates a single muscle - the superior oblique muscle of the eye (which operates through the pu ...
(type I) *The decussation of part of the
Oculomotor nerve The oculomotor nerve, also known as the third cranial nerve, cranial nerve III, or simply CN III, is a cranial nerve that enters the orbit through the superior orbital fissure and innervates extraocular muscles that enable most movements o ...
(type I) The optic chiasm of vertebrates involves the optic tract. The trochlear nerve is a
motor nerve A motor nerve, or efferent nerve, is a nerve that contains exclusively efferent nerve fibers and transmits motor signals from the central nervous system (CNS) to the effector organs (muscles and glands), as opposed to sensory nerves, which transf ...
that innervates one of the
muscles Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
that move the contralateral eye (i.e., the
superior oblique muscle The superior oblique muscle or obliquus oculi superior is a fusiform muscle originating in the upper, medial side of the orbit (anatomy), orbit (i.e. from beside the nose) which abducts, depresses and internally rotates the eye. It is the only e ...
). It emerges from the dorsal aspect of the ventral
midbrain The midbrain or mesencephalon is the uppermost portion of the brainstem connecting the diencephalon and cerebrum with the pons. It consists of the cerebral peduncles, tegmentum, and tectum. It is functionally associated with vision, hearing, mo ...
, leaves the brain on the dorsal side where it crosses to the opposite side. The
oculomotor nerve The oculomotor nerve, also known as the third cranial nerve, cranial nerve III, or simply CN III, is a cranial nerve that enters the orbit through the superior orbital fissure and innervates extraocular muscles that enable most movements o ...
originates from the third nerve
nucleus Nucleus (: nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucleu ...
at the level of the
superior colliculus In neuroanatomy, the superior colliculus () is a structure lying on the tectum, roof of the mammalian midbrain. In non-mammalian vertebrates, the Homology (biology), homologous structure is known as the optic tectum or optic lobe. The adjective f ...
(in non-mammalian vertebrates this is the
optic tectum In neuroanatomy, the superior colliculus () is a structure lying on the roof of the mammalian midbrain. In non-mammalian vertebrates, the homologous structure is known as the optic tectum or optic lobe. The adjective form '' tectal'' is commonly ...
) in the midbrain. The
rostral Rostral may refer to: Anatomy * Rostral (anatomical term), situated toward the oral or nasal region * Rostral bone, in ceratopsian dinosaurs * Rostral organ, of certain fish * Rostral scale The rostral scale, or rostral, in snakes and other sca ...
part of the nerve crosses the midline to merge with the part of the contralateral nerve that does not cross. Since the midline crossing occurs inside the brain, it is not strictly a chiasm but rather a decussation.


Structure

As stated above, very different kinds of nerve crossings are known as chiasm. The
optic chiasm In neuroanatomy, the optic chiasm, or optic chiasma (; , ), is the part of the brain where the optic nerves cross. It is located at the bottom of the brain immediately inferior to the hypothalamus. The optic chiasm is found in all vertebrates, ...
of vertebrates is the best known. The optic nerve runs from the
retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
towards the
ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
midline of the brain and crosses to the opposite side to continue as the optic tract which inserts to the
optic tectum In neuroanatomy, the superior colliculus () is a structure lying on the roof of the mammalian midbrain. In non-mammalian vertebrates, the homologous structure is known as the optic tectum or optic lobe. The adjective form '' tectal'' is commonly ...
(=superior colliculus) on the dorsal midbrain (as well as branching off to the
thalamus The thalamus (: thalami; from Greek language, Greek Wikt:θάλαμος, θάλαμος, "chamber") is a large mass of gray matter on the lateral wall of the third ventricle forming the wikt:dorsal, dorsal part of the diencephalon (a division of ...
in
amniotes Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the ...
).


Type I: Passing in the sagittal plane

In many vertebrates, the left-eye optic nerve crosses over the right-eye one, without blending.


Type II: Fusion in the sagittal plane

In
mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
and
birds Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
and other vertebrates with frontal eyes, the optic nerves do blend in the optic chiasm, and only part of the nerve fibres cross the midline. The drawings of Cajal suggest that the axons of the optic nerve may
branch A branch, also called a ramus in botany, is a stem that grows off from another stem, or when structures like veins in leaves are divided into smaller veins. History and etymology In Old English, there are numerous words for branch, includ ...
in the optic chiasm, and thus give off a branch both in the ipsi- and contralateral optic tract. Note, however, that such branching is not neural processing as occurs in a ganglion.


Type III: Systematic crossing of fibres inside a nerve

The optic tract of various clades of insects shows two chiasms, the first and second optic chiasm. In contrast to those in vertebrates, the insect chiasms do not cross the body midline. Rather, the first and second chiasm invert the anterior and posterior visual field. Since there are two chiasms, the retinotopic map is not affected. Cephalopods (
squids A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also called ''squi ...
and
octopuses An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed Mollusca, mollusc of the order (biology), order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, ...
) possess highly developed lens eyes. The optic tract of cephalopods, such as the squid ''
Loligo ''Loligo'' is a genus of squid and one of the most representative and widely distributed groups of myopsid squid. The genus was first described by Jean Baptiste Lamarck in 1798. However, the name had been used earlier than Lamarck ( Schneider, ...
'', chiasmates without midline crossing. This chiasm is distributed along the optic tract and effectively compensates the inversion of the image on the
retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
.


Type IV: Torsion or looping of a nerve

This type is usually not called chiasm. Such a looping occurs, for example, in the optic tract between the optic chiasm and the optic tectum. Another example is the
optic radiation In neuroanatomy, the optic radiation (also known as the geniculocalcarine tract, the geniculostriate pathway, and posterior thalamic radiation) are axons from the neurons in the lateral geniculate nucleus to the primary visual cortex. The optic ...
which rotates the retinal map on the visual cortex by 180° (see Figure 3).


Theories and evolution


Vertebrates

A number of theories have been proposed to explain the existence of the optic chiasm in vertebrates. The first is these theories was the Visual map theory by Ramón y Cajal. The
Axial twist theory The axial twist theory (wiktionary:also known as, a.k.a. axial twist ''hypothesis'') is a proposed scientific theory that has been proposed to explain a range of unusual aspects of the body plan of vertebrates (including humans). It states that th ...
also explains the chiasm of the trochlear nerve. The hypothesis of Cajal might be valid for the optic chiasm of cephalopods, although in a different manner, because Cajal designed his idea for a chiasm of type II but the cephalopod chiasm is of type III.


invertebrates

The lens eye inverts the visual image that is projected on the retina due to the
camera obscura A camera obscura (; ) is the natural phenomenon in which the rays of light passing through a aperture, small hole into a dark space form an image where they strike a surface, resulting in an inverted (upside down) and reversed (left to right) ...
effect. The chiasm in the optic tract of cephalopods corrects this inversion. In insects, the optic chiasms seem to have evolved gradually, since primitive groups have no chiasm, whereas later evolved groups have one or two optic chiasms along the optic lobe.


Exceptions

In
jawless vertebrates Agnatha (; ) or jawless fish is a paraphyletic infraphylum of animals in the subphylum Vertebrata of the phylum Chordata, characterized by the lack of jaws. The group consists of both extant taxon, living (Cyclostomi, cyclostomes such as hagfish ...
(
hagfish Hagfish, of the Class (biology), class Myxini (also known as Hyperotreti) and Order (biology), order Myxiniformes , are eel-shaped Agnatha, jawless fish (occasionally called slime eels). Hagfish are the only known living Animal, animals that h ...
and
lamprey Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
), the optic tracts do cross in the midline, but only after entering the ventral side of the central nervous system. After crossing the tracts insert on the dorsal optic tectum as in all other vertebrates. Therefore, given the obvious and undisputed homology, the optic chiasm is called chiasm also in these clades, even though the crossing is technically a decussation.


See also

* Definition of types of crossings *
Contralateral brain The contralateral organization of the forebrain (Latin: contra‚ against; latus‚ side; lateral‚ sided) is the property that the Cerebral hemisphere, hemispheres of the cerebrum and the thalamus represent mainly the contralateral side of the b ...
*
Optic chiasm In neuroanatomy, the optic chiasm, or optic chiasma (; , ), is the part of the brain where the optic nerves cross. It is located at the bottom of the brain immediately inferior to the hypothalamus. The optic chiasm is found in all vertebrates, ...
* Chiasmal syndromes *
Insect morphology Insect morphology is the study and description of the morphology (biology), physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical f ...
*
Chiasmatic groove The chiasmatic groove (chiasmatic sulcus, optic groove, prechiasmatic sulcus) is a transverse groove upon the superior aspect of the body of sphenoid bone within the middle cranial fossa. It is bounded anteriorly by the sphenoidal limbus (a variab ...


References


External links

* (human) * (non-human) {{Authority control Neuroanatomy Optic nerve Nerves Peripheral nervous system Soft tissue