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 optic chiasm, or optic chiasma (; , ), is the part of the

brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...

where the optic nerves cross. It is located at the bottom of the brain immediately inferior to the

hypothalamus The hypothalamus (: hypothalami; ) is a small part of the vertebrate brain that contains a number of nucleus (neuroanatomy), nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrin ...

. The optic chiasm is found in all

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 ...

, although in cyclostomes (

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 ...

s and

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 ...

es), it is located within the brain. This article is about the optic chiasm of vertebrates, which is the best known nerve chiasm, but not every chiasm denotes a crossing of the body midline (e.g., in some

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 ...

, see Chiasm (anatomy)). A midline crossing of nerves inside the brain is called 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 ...

(see Definition of types of crossings).

Structure

In all vertebrates, the optic nerves of the left and the right eye meet in the body midline, ventral to the brain. In many vertebrates the left optic nerve crosses over the right one without fusing with it. In vertebrates with a large overlap of the visual fields of the two eyes, i.e., most mammals and birds, but also

amphibians Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

s such as chameleons, the two optic nerves merge in the optic chiasm. In such a merged optic chiasm, part of the nerve fibres do not cross the midline, but continue towards 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 ...

of the ipsilateral side. By this partial decussation, the part of the

visual field The visual field is "that portion of space in which objects are visible at the same moment during steady fixation of the gaze in one direction"; in ophthalmology and neurology the emphasis is mostly on the structure inside the visual field and it i ...

that is covered by both eyes is fused so that the processing of binocular

depth perception Depth perception is the ability to perceive distance to objects in the world using the visual system and visual perception. It is a major factor in perceiving the world in three dimensions. Depth sensation is the corresponding term for non-hum ...

stereopsis Binocular vision is seeing with two eyes, which increases the size of the Visual field, visual field. If the visual fields of the two eyes overlap, binocular #Depth, depth can be seen. This allows objects to be recognized more quickly, camouflage ...

is enabled (see Figure 2). In the case of such partial decussation, the optic nerve fibres on the medial sides of each

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 ...

(which correspond to the lateral side of each visual hemifield, because the image is inverted) cross over to the opposite side of the body midline. The inferonasal retina are related to the anterior portion of the optic chiasm whereas superonasal retinal fibers are related to the posterior portion of the optic chiasm. The partial crossing over of optic nerve fibres at the optic chiasm allows the visual cortex to receive the same hemispheric

from both eyes. Superimposing and processing these monocular visual signals allow the visual cortex to generate binocular and

stereoscopic Stereoscopy, also called stereoscopics or stereo imaging, is a technique for creating or enhancing the illusion of depth in an image by means of stereopsis for binocular vision. The word ''stereoscopy'' derives . Any stereoscopic image is ...

vision. The net result is that the right cerebral hemisphere processes left visual hemifield, and the left cerebral hemisphere processes the right visual hemifield. Beyond the optic chiasm, with crossed and uncrossed fibers, the optic nerves are called optic tracts. The optic tract inserts on the optic tectum (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 ...

known as

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 ...

) of the

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 ...

. In mammals they also branch off to the lateral geniculate body of 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 turn giving them to the occipital cortex of the

cerebrum The cerebrum (: cerebra), telencephalon or endbrain is the largest part of the brain, containing the cerebral cortex (of the two cerebral hemispheres) as well as several subcortical structures, including the hippocampus, basal ganglia, and olfac ...

Arterial supply

The optic chiasma receives its arterial supply from the anterior cerebral arteries, and from branches of the

internal carotid artery The internal carotid artery is an artery in the neck which supplies the anterior cerebral artery, anterior and middle cerebral artery, middle cerebral circulation. In human anatomy, the internal and external carotid artery, external carotid ari ...

which ascend along the

pituitary stalk The pituitary stalk, also known as the infundibular stalk, infundibulum, or Fenderson's funnel, is the connection between the hypothalamus and the posterior pituitary, the posterior lobe of the pituitary gland. The floor of the third ventricle is ...

(the latter supplying the midline portion of the chiasma).

Development in mammals

During

development Development or developing may refer to: Arts *Development (music), the process by which thematic material is reshaped * Photographic development *Filmmaking, development phase, including finance and budgeting * Development hell, when a proje ...

, the crossing of the optic nerves is guided primarily by cues such as netrin, slit,

semaphorin Semaphorins are a class of secreted and membrane proteins that were originally identified as axonal growth cone guidance molecules. They primarily act as short-range inhibitory signals and signal through multimeric receptor (biochemistry), recepto ...

and ephrin; and by

morphogens A morphogen is a substance whose non-uniform distribution governs the Natural patterns, pattern of tissue development in the process of morphogenesis or pattern formation, one of the core processes of developmental biology, establishing positio ...

such as

sonic hedgehog Sonic hedgehog protein (SHH) is a major signaling molecule of embryonic development in humans and animals, encoded by the ''SHH'' gene. This signaling molecule is key in regulating embryonic morphogenesis in all animals. SHH controls organoge ...

(Shh) and Wnt. This navigation is mediated by the neuronal

growth cone A growth cone is a large actin-supported extension of a developing or regenerating neurite seeking its synaptic target. It is the growth cone that drives axon growth. Their existence was originally proposed by Spanish histologist Santiago ...

, a structure that responds to the cues by

ligand In coordination chemistry, a ligand is an ion or molecule with a functional group that binds to a central metal atom to form a coordination complex. The bonding with the metal generally involves formal donation of one or more of the ligand's el ...

receptor Receptor may refer to: * Sensory receptor, in physiology, any neurite structure that, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and respond ...

signalling systems that activate downstream pathways inducing changes in the

cytoskeleton The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compos ...

Retinal ganglion cell A retinal ganglion cell (RGC) is a type of neuron located near the inner surface (the ganglion cell layer) of the retina of the eye. It receives visual information from photoreceptor cell, photoreceptors via two intermediate neuron types: Bipolar ...

(RGC) axons leaving the eye through the optic nerve are blocked from exiting the developing pathway by Slit2 and Sema5A inhibition, expressed bordering the optic nerve pathway. Ssh expressed at 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 ...

midline inhibits crossing prior to the chiasm, where it is downregulated. The organization of RGC axons changes from

retinotopic Retinotopy () is the mapping of visual input from the retina to neurons, particularly those neurons within the Two-streams hypothesis, visual stream. For clarity, 'retinotopy' can be replaced with 'retinal mapping', and 'retinotopic' with 'retin ...

to a flat sheet-like orientation as they approach the chiasm site. Most RGC

axon An axon (from Greek ἄξων ''áxōn'', axis) or nerve fiber (or nerve fibre: see American and British English spelling differences#-re, -er, spelling differences) is a long, slender cellular extensions, projection of a nerve cell, or neuron, ...

s cross the midline at 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 ...

diencephalon In the human brain, the diencephalon (or interbrain) is a division of the forebrain (embryonic ''prosencephalon''). It is situated between the telencephalon and the midbrain (embryonic ''mesencephalon''). The diencephalon has also been known as t ...

and continue to the

contralateral 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 ...

. The number of axons that do not cross the midline and project ipsilaterally depends on the degree of binocular vision of the animal (3% in mice and 45% in humans do not cross). Ephrin-B2 is expressed at the chiasm midline by radial glia and acts as a repulsive signal to axons originating from the ventrotemporal retina expressing EphB1 receptor protein, giving rise to the ipsilateral, or uncrossed, projection. RGC axons that do cross at the optic chiasm are guided by the vascular endothelial growth factor, VEGF-A, expressed at the midline, which signals through the receptor Neuropilin-1 (NRP1) expressed on RGC axons. Chiasm crossing is also promoted by Nr-CAM (Ng-CAM-related cell adhesion molecule) and

Semaphorin Semaphorins are a class of secreted and membrane proteins that were originally identified as axonal growth cone guidance molecules. They primarily act as short-range inhibitory signals and signal through multimeric receptor (biochemistry), recepto ...

6D (Sema6D) expressed at the midline, which form a complex that signals to Nr-CAM/

Plexin A plexin is a protein which acts as a receptor for semaphorin family signaling proteins. It is classically known for its expression on the surface of axon growth cones and involvement in signal transduction to steer axon growth away from the sour ...

-A1 receptors on crossing RGC axons.

Other animals

Mammals

Since all vertebrates, even the earliest fossils and modern jawless ones, possess an optic chiasm, it is not known how it evolved. A number of theories have been proposed for the function of the optic chiasm in vertebrates (see

theories A theory is a systematic and rational form of abstract thinking about a phenomenon, or the conclusions derived from such thinking. It involves contemplative and logical reasoning, often supported by processes such as observation, experimentation, ...

). According to the Axial Twist theory the optic chiasm develops as a consequence of a twist in the early

embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...

. In Siamese cats with certain genotypes of the

albino Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and reddish pink or blue eyes. Individuals with the condition are referred to as albinos. Varied use and interpretation of ...

gene, the wiring is disrupted, with more of the nerve-crossing than normal. Since siamese cats, like

tigers, also tend to cross their eyes (

strabismus Strabismus is an eye disorder in which the eyes do not properly align with each other when looking at an object. The eye that is pointed at an object can alternate. The condition may be present occasionally or constantly. If present during a ...

), it has been proposed that this behavior might compensate the abnormal amount of

Cephalopods and insects

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 ...

and

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 ...

the optic tracts do not cross the body midline, so each side of the brain processes the ipsilateral eye.

History

The crossing of nerve fibres, and the impact on vision that this had, was probably first identified by Persian physician "Esmail Jorjani", who appears to be Zayn al-Din Gorgani (1042–1137).

Additional images

File:Gray722.svg, Scheme showing central connections of the

optic nerve In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual system, visual information from the retina to the brain. In humans, the optic nerve i ...

s and optic tracts. File:Gray724.png, Brain seen from below, with the optic chiasm seen in yellow in the centre. File:Optical-transformations.png, Transformations of the visual field toward the visual map on the primary visual cortex. File:Human brainstem anterior view 2 description.JPG, Brain and brainstem seen from below File:Slide13qq.JPG, Left hemisphere of the brain seen in a cadaveric specimen from the side, with the optic chiasm labelled. File:Slide2Dsa.JPG, Cerebrum, inferior view, deep dissection. File:Optic chiasm development.jpg, Guidance of axon crossing and non-crossing during development.

References

External links

* {{Authority control Cerebrum Visual system Optic nerve