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, defin ...
, the superior colliculus () is a structure lying on the
roof
A roof ( : roofs or rooves) is the top covering of a building, including all materials and constructions necessary to support it on the walls of the building or on uprights, providing protection against rain, snow, sunlight, extremes of temper ...
of the mammalian
midbrain
The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', " ...
.
In non-mammalian
vertebrates
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
, the
homologous structure is known as the optic tectum, or optic lobe.
The adjective form ''
tectal
The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', "m ...
'' is commonly used for both structures.
In mammals, the superior colliculus forms a major component of the midbrain. It is a paired structure and together with the paired
inferior colliculi
The inferior colliculus (IC) (Latin for ''lower hill'') is the principal midbrain nucleus of the auditory pathway and receives input from several peripheral brainstem nuclei in the auditory pathway, as well as inputs from the auditory cortex. Th ...
forms the
corpora quadrigemina
Corpus is Latin for "body". It may refer to:
Linguistics
* Text corpus, in linguistics, a large and structured set of texts
* Speech corpus, in linguistics, a large set of speech audio files
* Corpus linguistics, a branch of linguistics
Music
* ...
. The superior colliculus is a layered structure, with a pattern that is similar to all mammals.
The layers can be grouped into the superficial layers (
stratum opticum
The retinal nerve fiber layer (RNFL) or nerve fiber layer, stratum opticum, is formed by the expansion of the fibers of the optic nerve; it is thickest near the optic disc, gradually diminishing toward the ora serrata.
As the nerve fibers pass t ...
and above) and the deeper remaining layers. Neurons in the superficial layers receive direct input from the
retina
The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
and respond almost exclusively to visual stimuli. Many neurons in the deeper layers also respond to other modalities, and some respond to stimuli in multiple modalities. The deeper layers also contain a population of motor-related neurons, capable of activating
eye movement
Eye movement includes the voluntary or involuntary movement of the eyes. Eye movements are used by a number of organisms (e.g. primates, rodents, flies, birds, fish, cats, crabs, octopus) to fixate, inspect and track visual objects of interests. ...
s as well as other responses. In other vertebrates the number of layers in the homologous optic tectum varies.
The general function of the tectal system is to direct behavioral responses toward specific points in body-centered space. Each layer contains a
topographic map
In modern mapping, a topographic map or topographic sheet is a type of map characterized by large- scale detail and quantitative representation of relief features, usually using contour lines (connecting points of equal elevation), but historic ...
of the surrounding world in
retinotopic
Retinotopy (from Greek τόπος, place) is the mapping of visual input from the retina to neurons, particularly those neurons within the visual stream. For clarity, 'retinotopy' can be replaced with 'retinal mapping', and 'retinotopic' with 'r ...
coordinates, and activation of neurons at a particular point in the map evokes a response directed toward the corresponding point in space. In primates, the superior colliculus has been studied mainly with respect to its role in directing eye movements. Visual input from the retina, or "command" input from the cerebral cortex, create a "bump" of activity in the tectal map, which, if strong enough, induces a
saccadic eye movement
A saccade ( , French for ''jerk'') is a quick, simultaneous movement of both eyes between two or more phases of fixation in the same direction.Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishin ...
. Even in primates, however, the superior colliculus is also involved in generating spatially directed head turns, arm-reaching movements, and shifts in attention that do not involve any overt movements.
In other species, the superior colliculus is involved in a wide range of responses, including whole-body turns in walking rats. In mammals, and especially primates, the massive expansion of the cerebral cortex reduces the superior colliculus to a much smaller fraction of the whole brain. It remains nonetheless important in terms of function as the primary integrating center for eye movements.
In non-mammalian species the optic tectum is involved in many responses including swimming in fish, flying in birds, tongue-strikes toward prey in frogs, and fang-strikes in snakes. In some species, including fish and birds, the optic tectum, also known as the optic lobe, is one of the largest components of the brain.
''Note on terminology'': This article follows terminology established in the literature, using the term "superior colliculus" when discussing mammals and "optic tectum" when discussing either specific non-mammalian species or vertebrates in general.
Structure
![Gray719](https://upload.wikimedia.org/wikipedia/commons/0/0b/Gray719.png)
The superior colliculus is a synaptic layered structure.
The two superior colliculi sit below the
thalamus
The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
and surround the
pineal gland
The pineal gland, conarium, or epiphysis cerebri, is a small endocrine gland in the brain of most vertebrates. The pineal gland produces melatonin, a serotonin-derived hormone which modulates sleep, sleep patterns in both circadian rhythm, circ ...
in the
mammalian
Mammals () are a group of vertebrate animals constituting the class (biology), class Mammalia (), characterized by the presence of mammary glands which in Female#Mammalian female, females produce milk for feeding (nursing) their young, a ...
midbrain
The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', " ...
. It comprises the
dorsal
Dorsal (from Latin ''dorsum'' ‘back’) may refer to:
* Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism
* Dorsal, positioned on top of an aircraft's fuselage
* Dorsal co ...
aspect of the
midbrain
The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', " ...
, posterior to the
periaqueductal gray
The periaqueductal gray (PAG, also known as the central gray) is a brain region that plays a critical role in autonomic function, motivated behavior and behavioural responses to threatening stimuli. PAG is also the primary control center for d ...
and immediately superior to the
inferior colliculus
The inferior colliculus (IC) (Latin for ''lower hill'') is the principal midbrain nucleus of the auditory pathway and receives input from several peripheral brainstem nuclei in the auditory pathway, as well as inputs from the auditory cortex. The ...
. The inferior and superior colliculi are known collectively as the
corpora quadrigemina
Corpus is Latin for "body". It may refer to:
Linguistics
* Text corpus, in linguistics, a large and structured set of texts
* Speech corpus, in linguistics, a large set of speech audio files
* Corpus linguistics, a branch of linguistics
Music
* ...
(Latin for ''quadruplet bodies''). The superior colliculi are larger than the inferior colliculi, though the inferior colliculi are more prominent.
The brachium of superior colliculus (or superior brachium) is a branch that extends laterally from the superior colliculus, and, passing to the
thalamus
The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
between the
pulvinar and the
medial geniculate nuclei
The medial geniculate nucleus (MGN) or medial geniculate body (MGB) is part of the auditory thalamus and represents the thalamic relay between the inferior colliculus (IC) and the auditory cortex (AC). It is made up of a number of sub-nuclei that ...
, is partly continued into an eminence called the
lateral geniculate nucleus
In neuroanatomy, the lateral geniculate nucleus (LGN; also called the lateral geniculate body or lateral geniculate complex) is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, ventral projec ...
, and partly into 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 supe ...
.
The superior colliculus is associated with a nearby structure called the parabigeminal nucleus, often referred to as its satellite. In the optic tectum this nearby structure is known as the
nucleus isthmi
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 ...
.
Neural circuitry
The microstructure of the superior colliculus and of the optic tectum, varies across species. As a general rule, there is always a clear distinction between superficial layers, which receive input primarily from the visual system and show primarily visual responses, and deeper layers, which receive many types of input and project to numerous motor-related brain areas. The distinction between these two zones is so clear and consistent that some anatomists have suggested that they should be considered separate brain structures.
In mammals, neuroanatomists conventionally identify seven layers The top three layers are called ''superficial'':
*Lamina I or SZ, the ''stratum zonale'', is a thin layer consisting of small myelinated axons together with marginal and horizontal cells.
*Lamina II or SGS, the ''stratum griseum superficiale'' ("superficial gray layer"), contains many neurons of various shapes and sizes.
*Lamina III or SO, the ''stratum opticum'' ("optic layer"), consists mainly of axons coming from 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 supe ...
.
Next come two ''intermediate layers'':
*Lamina IV or SGI, the ''stratum griseum intermedium'' ("intermediate gray layer"), is the thickest layer, and is filled with many neurons of many sizes. This layer is often as thick as all the other layers together. It is often subdivided into "upper" and "lower" parts.
*Lamina V or SAI, the ''stratum album intermedium'' ("intermediate white layer"), consists mainly of fibers from various sources.
Finally come the two ''deep layers'':
*Lamina VI or SGP, the ''stratum griseum profundum'' ("deep gray layer"), consists of loosely packed neurons and myelinated fibers.
*Lamina VII or SAP, the ''stratum album profundum'' ("deep white layer"), lying directly above the
periaqueductal gray
The periaqueductal gray (PAG, also known as the central gray) is a brain region that plays a critical role in autonomic function, motivated behavior and behavioural responses to threatening stimuli. PAG is also the primary control center for d ...
, consists entirely of fibers.
The superficial layers receive input mainly from the retina, vision-related areas of the cerebral cortex, and two tectal-related structures called the
pretectum
In neuroanatomy, the pretectal area, or pretectum, is a midbrain structure composed of seven nuclei and comprises part of the subcortical visual system. Through reciprocal bilateral projections from the retina, it is involved primarily in mediati ...
and ''parabigeminal nucleus''. The retinal input encompasses the entire superficial zone, and is bilateral, although the contralateral portion is more extensive. The cortical input comes most heavily from the
primary visual cortex
The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalamus and ...
(area 17, V1), the secondary visual cortex (areas
18 and
19), and the
frontal eye fields
The frontal eye fields (FEF) are a region located in the frontal cortex, more specifically in Brodmann area 8 or BA8, of the primate brain. In humans, it can be more accurately said to lie in a region around the intersection of the middle fronta ...
. The parabigeminal nucleus plays a very important role in tectal function that is described below.
In contrast to the vision-dominated inputs to the superficial layers, the intermediate and deep layers receive inputs from a very diverse set of sensory and motor structures. Most areas of the cerebral cortex project to these layers, although the input from "association" areas tends to be heavier than the input from primary sensory or motor areas. However, the cortical areas involved, and the strength of their relative projections differs across species. Another important input comes from the
substantia nigra
The substantia nigra (SN) is a basal ganglia structure located in the midbrain that plays an important role in reward and movement. ''Substantia nigra'' is Latin for "black substance", reflecting the fact that parts of the substantia nigra app ...
,
pars reticulata
The pars reticulata (SNpr) is a portion of the substantia nigra and is located lateral to the pars compacta. Most of the neurons that project out of the pars reticulata are inhibitory GABAergic neurons (i.e., these neurons release GABA, which is ...
, a component of the
basal ganglia
The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an extern ...
. This projection uses the inhibitory neurotransmitter
GABA, and is thought to exert a "gating" effect on the superior colliculus. The intermediate and deep layers also receive input from the
spinal trigeminal nucleus
The spinal trigeminal nucleus is a nucleus in the medulla that receives information about deep/crude touch, pain, and temperature from the ipsilateral face. In addition to the trigeminal nerve (CN V), the facial (CN VII), glossopharyngeal (CN IX), ...
, which conveys somatosensory information from the face, as well as the
hypothalamus
The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamu ...
,
zona incerta
The zona incerta (ZI) is a horizontally elongated region of gray matter in the subthalamus below the thalamus. Its connections project extensively over the brain from the cerebral cortex down into the spinal cord.
Its function is unknown, though ...
,
thalamus
The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
, and
inferior colliculus
The inferior colliculus (IC) (Latin for ''lower hill'') is the principal midbrain nucleus of the auditory pathway and receives input from several peripheral brainstem nuclei in the auditory pathway, as well as inputs from the auditory cortex. The ...
.
In addition to their distinctive inputs, the superficial and deep zones of the superior colliculus also have distinctive outputs. One of the most important outputs goes to the
pulvinar and lateral intermediate areas of the thalamus, which in turn project to areas of the cerebral cortex that are involved in controlling eye movements. There are also projections from the superficial zone to the pretectal nuclei,
lateral geniculate nucleus
In neuroanatomy, the lateral geniculate nucleus (LGN; also called the lateral geniculate body or lateral geniculate complex) is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, ventral projec ...
of the thalamus, and the parabigeminal nucleus. The projections from the deeper layers are more extensive. There are two large descending pathways, traveling to the brainstem and spinal cord, and numerous ascending projections to a variety of sensory and motor centers, including several that are involved in generating eye movements.
Both colliculi also have descending projections to the
paramedian pontine reticular formation
The paramedian pontine reticular formation, also known as PPRF or paraabducens nucleus, is part of the pontine reticular formation, a brain region without clearly defined borders in the center of the pons. It is involved in the coordination of eye ...
and spinal cord, and thus can be involved in responses to stimuli faster than cortical processing would allow.
Mosaic structure
On detailed examination the collicular layers are actually not smooth sheets, but divided into a honeycomb arrangement of discrete columns.
[ Chavalier & Mana, 2000] The clearest indication of columnar structure comes from the cholinergic inputs arising from the parabigeminal nucleus, whose terminals form evenly spaced clusters that extend from top to bottom of the
tectum
The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', "m ...
.
[ Illing, 1996] Several other neurochemical markers including calretinin, parvalbumin, GAP-43, and NMDA receptors, and connections with numerous other brain structures in the brainstem and diencephalon, also show a corresponding inhomogeneity. The total number of columns has been estimated at around 100.
The functional significance of this columnar architecture is not clear, but it is interesting that recent evidence has implicated the cholinergic inputs as part of a recurrent circuit producing winner-take-all dynamics within the tectum, as described in more detail below.
All species that have been examined — including mammals and non-mammals — show compartmentalization, but there are some systematic differences in the details of the arrangement.
In species with a streak-type retina (mainly species with laterally placed eyes, such as rabbits and deer), the compartments cover the full extent of the SC. In species with a centrally placed fovea, however, the compartmentalization breaks down in the front (rostral) part of the SC. This portion of the SC contains many "fixation" neurons that fire continually while the eyes remain fixed in a constant position.
Function
The history of investigation of the optic tectum has been marked by several large shifts in opinion. Before about 1970, most studies involved non-mammals — fish, frogs, birds — that is, species in which the optic tectum is the dominant structure that receives input from the eyes. The general view then was that the optic tectum, in these species, is the main visual center in the non-mammalian brain, and, as a consequence, is involved in a wide variety of behaviors. From the 1970s to 1990s, however, neural recordings from mammals, mostly monkeys, focused primarily on the role of the superior colliculus in controlling eye movements. This line of investigation came to dominate the literature to such a degree that the majority opinion was that eye-movement control is the only important function in mammals, a view still reflected in many current textbooks.
In the late 1990s, however, experiments using animals whose heads were free to move showed clearly that the SC actually produces ''gaze shifts'', usually composed of combined head and eye movements, rather than eye movements ''per se''. This discovery reawakened interest in the full breadth of functions of the superior colliculus, and led to studies of
multisensory integration
Multisensory integration, also known as multimodal integration, is the study of how information from the different sensory modalities (such as sight, sound, touch, smell, self-motion, and taste) may be integrated by the nervous system. A coherent r ...
in a variety of species and situations. Nevertheless, the role of the SC in controlling eye movements is understood in much greater depth than any other function.
Behavioral studies have shown that the SC is not needed for object recognition, but plays a critical role in the ability to direct behaviors toward specific objects, and can support this ability even in the absence of the cerebral cortex. Thus, cats with major damage to the visual cortex cannot recognize objects, but may still be able to follow and orient toward moving stimuli, although more slowly than usual. If one half of the SC is removed, however, the cats will circle constantly toward the side of the lesion, and orient compulsively toward objects located there, but fail to orient at all toward objects located in the opposite hemifield. These deficits diminish over time but never disappear.
Eye movements
In primates,
eye movements
Eye movement includes the voluntary or involuntary movement of the eyes. Eye movements are used by a number of organisms (e.g. primates, rodents, flies, birds, fish, cats, crabs, octopus) to fixate, inspect and track visual objects of inte ...
can be divided into several types:
fixation, in which the eyes are directed toward a motionless object, with eye movements only to compensate for movements of the head;
smooth pursuit
In the scientific study of vision, smooth pursuit describes a type of eye movement in which the eyes remain fixated on a moving object. It is one of two ways that visual animals can voluntarily shift gaze, the other being saccadic eye movemen ...
, in which the eyes move steadily to track a moving object;
saccade
A saccade ( , French for ''jerk'') is a quick, simultaneous movement of both eyes between two or more phases of fixation in the same direction.Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishi ...
s, in which the eyes move very rapidly from one location to another; and
vergence
A vergence is the simultaneous movement of both eyes in opposite directions to obtain or maintain single binocular vision.
When a creature with binocular vision looks at an object, the eyes must rotate around a vertical axis so that the proje ...
, in which the eyes move simultaneously in opposite directions to obtain or maintain single binocular vision. The superior colliculus is involved in all of these, but its role in saccades has been studied most intensively.
Each of the two colliculi — one on each side of the brain — contains a two-dimensional map representing half of the visual field. The
fovea
Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure.
Human anatomy
*Fovea centralis of the retina
* Fovea buccalis or Dimple
* Fovea of the femoral head
* Trochlear fovea of the fr ...
— the region of maximum sensitivity — is represented at the front edge of the map, and the periphery at the back edge. Eye movements are evoked by activity in the deep layers of the SC. During fixation, neurons near the front edge — the foveal zone — are tonically active. During smooth pursuit, neurons a small distance from the front edge are activated, leading to small eye movements. For saccades, neurons are activated in a region that represents the point to which the saccade will be directed. Just prior to a saccade, activity rapidly builds up at the target location and decreases in other parts of the SC. The coding is rather broad, so that for any given saccade the activity profile forms a "hill" that encompasses a substantial fraction of the collicular map: The location of the peak of this "hill" represents the saccade target.
The SC encodes the target of a gaze shift, but it does not seem to specify the precise movements needed to get there. The decomposition of a gaze shift into head and eye movements and the precise trajectory of the eye during a saccade depend on integration of collicular and non-collicular signals by downstream motor areas, in ways that are not yet well understood. Regardless of how the movement is evoked or performed, the SC encodes it in "retinotopic" coordinates: that is, the location of the SC 'hill" corresponds to a fixed location on the retina. This seems to contradict the observation that stimulation of a single point on the SC can result in different gaze shift directions, depending on initial eye orientation. However, it has been shown that this is because the retinal location of a stimulus is a non-linear function of target location, eye orientation, and the spherical geometry of the eye.
There has been some controversy about whether the SC merely commands eye movements, and leaves the execution to other structures, or whether it actively participates in the performance of a saccade. In 1991, Munoz et al., on the basis of data they collected, argued that, during a saccade, the "hill" of activity in the SC moves gradually, to reflect the changing offset of the eye from the target location while the saccade is progressing. At present, the predominant view is that, although the "hill" does shift slightly during a saccade, it does not shift in the steady and proportionate way that the "moving hill" hypothesis predicts. However, moving hills may play another role in the superior colliculus; more recent experiments have demonstrated a continuously moving hill of visual memory activity when the eyes move slowly while a separate saccade target is retained.
The output from the motor sector of the SC goes to a set of midbrain and brainstem nuclei, which transform the "place" code used by the SC into the "rate" code used by oculomotor neurons. Eye movements are generated by six muscles, arranged in three orthogonally-aligned pairs. Thus, at the level of the final common path, eye movements are encoded in essentially a Cartesian coordinate system.
Although the SC receives a strong input directly from the retina, in primates it is largely under the control of the cerebral cortex, which contains several areas that are involved in determining eye movements. The
frontal eye fields
The frontal eye fields (FEF) are a region located in the frontal cortex, more specifically in Brodmann area 8 or BA8, of the primate brain. In humans, it can be more accurately said to lie in a region around the intersection of the middle fronta ...
, a portion of the motor cortex, are involved in triggering intentional saccades, and an adjoining area, the supplementary eye fields, are involved in organizing groups of saccades into sequences. The parietal eye fields, farther back in the brain, are involved mainly in reflexive saccades, made in response to changes in the view. Recent evidence suggests that the
primary visual cortex
The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalamus and ...
(V1) guides reflexive eye movements, according to
V1 Saliency Hypothesis
The V1 Saliency Hypothesis, or V1SH (pronounced‘vish’) is a theory about V1, the primary visual cortex (V1). It proposes that the V1 in primates creates a saliency map of the visual field to guide visual attention or gaze shifts exogenously.
...
, using a bottom-up saliency map of the visual field generated in V1 from external visual inputs.
The SC only receives
visual
The visual system comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight (the ...
inputs in its superficial layers, whereas the deeper layers of the colliculus receive also auditory and somatosensory inputs and are connected to many sensorimotor areas of the brain. The colliculus as a whole is thought to help orient the head and eyes toward something seen and heard.
[ Kustov & Robinson, 1996]
The superior colliculus also receives auditory information from the inferior colliculus. This auditory information is integrated with the visual information already present to produce the
ventriloquism effect
In psychology, visual capture is the dominance of vision over other sense modalities in creating a percept.Grünwald, Martin. ''Human haptic perception''. Birkhauser, 2008. 657. Print. In this process, the visual senses influence the other parts o ...
.
Distractibility
As well as being related to eye movements, the SC appears to have an important role to play in the circuitry underpinning distractibility. Heightened distractibility occurs in normal aging and is also a central feature in a number of medical conditions, including
attention deficit hyperactivity disorder
Attention deficit hyperactivity disorder (ADHD) is a neurodevelopmental disorder characterised by excessive amounts of inattention, hyperactivity, and impulsivity that are pervasive, impairing in multiple contexts, and otherwise age-inap ...
(ADHD). Research has shown that lesions to the SC in a number of species can result in heightened distractibility and, in humans, removing the inhibitory control on the SC from the pre-frontal cortex, therefore increasing activity in the area, also increases distractibility. Research in an animal model of ADHD, the spontaneously hypertensive rat, also shows altered collicular-dependent behaviours
and physiology.
Furthermore, amphetamine (a mainstay treatment for ADHD) also suppresses activity in the colliculus in healthy animals.
Other animals
Other mammals
Primates
It is usually accepted that the
primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
superior colliculus is unique among
mammals
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
, in that it does not contain a complete map of the visual field seen by the contralateral eye. Instead, like the
visual cortex
The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalamus and ...
and
lateral geniculate nucleus
In neuroanatomy, the lateral geniculate nucleus (LGN; also called the lateral geniculate body or lateral geniculate complex) is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, ventral projec ...
, each colliculus represents only the contralateral half of the
visual field
The visual field is the "spatial array of visual sensations available to observation in introspectionist psychological experiments". Or simply, visual field can be defined as the entire area that can be seen when an eye is fixed straight at a point ...
, up to the midline, and excludes a representation of the ipsilateral half. This functional characteristic is explained by the absence, in primates, of anatomical connections between the
retinal ganglion cells
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 photoreceptors via two intermediate neuron types: bipolar cells and retina ...
in the temporal half of the
retina
The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
and the contralateral superior colliculus. In other mammals, the retinal ganglion cells throughout the contralateral retina project to the contralateral colliculus. This distinction between primates and non-primates has been one of the key lines of evidence in support of the
flying primates theory In evolutionary biology, the flying primate hypothesis is that megabats, a subgroup of Chiroptera (also known as flying foxes), form an evolutionary sister group of primates. The hypothesis began with Carl Linnaeus in 1758, and was again advanced by ...
proposed by Australian neuroscientist
Jack Pettigrew
John Douglas "Jack" Pettigrew (2 October 1943''Who's Who in Australia 2013'', Crown Content, 2012. - 7 May 2019) was an Australian neuroscientist. He was Emeritus Professor of Physiology and Director of the Vision, Touch and Hearing Research ...
in 1986, after he discovered that flying foxes (
megabat
Megabats constitute the family Pteropodidae of the order Chiroptera (bats). They are also called fruit bats, Old World fruit bats, or—especially the genera ''Acerodon'' and ''Pteropus''—flying foxes. They are the only member of the su ...
s) resemble primates in terms of the pattern of anatomical connections between the retina and superior colliculus.
Cats
In the cat the superior colliculus projects through the
reticular formation
The reticular formation is a set of interconnected nuclei that are located throughout the brainstem. It is not anatomically well defined, because it includes neurons located in different parts of the brain. The neurons of the reticular formation ...
and interacts with motor neurons in the
brainstem
The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is cont ...
.
Bats
Bat
Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s are not, in fact, blind, but they depend much more on echolocation than vision for navigation and prey capture. They obtain information about the surrounding world by emitting sonar chirps and then listening for the echoes. Their brains are highly specialized for this process, and some of these specializations appear in the superior colliculus. In bats, the retinal projection occupies only a thin zone just beneath the surface, but there are extensive inputs from auditory areas, and outputs to motor areas capable of orienting the ears, head, or body. Echoes coming from different directions activate neurons at different locations in the collicular layers,
[ Valentine & Moss, 1997] and activation of collicular neurons influences the chirps that the bats emit. Thus, there is a strong case that the superior colliculus performs the same sorts of functions for the auditory-guided behaviors of bats that it performs for the visual-guided behaviors of other species.
Bats are usually classified into two main groups:
Microchiroptera (the most numerous, and commonly found throughout the world), and
Megachiroptera (fruit bats, found in Asia, Africa and Australasia). With one exception,
Megabats
Megabats constitute the family Pteropodidae of the order Chiroptera ( bats). They are also called fruit bats, Old World fruit bats, or—especially the genera ''Acerodon'' and ''Pteropus''—flying foxes. They are the only member of the sup ...
do not echolocate, and rely on a developed sense of vision to navigate. The visual
receptive fields
The receptive field, or sensory space, is a delimited medium where some Stimulus (physiology), physiological stimuli can evoke a Sensory neuron, sensory neuronal response in specific organisms.
Complexity of the receptive field ranges from the un ...
of neurons in the superior colliculus in these animals form a precise map of the
retina
The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
, similar to that found in
cat
The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s and
primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s.
Rodents
The superior colliculus in rodents have been hypothesized to mediate sensory-guided approach and avoidance behaviors. Studies employing circuit analysis tools on mouse superior colliculus have revealed several important functions.
In a series of studies, researchers have identified a set of Ying-Yang circuit modules in the superior colliculus to initiate prey capture and predator avoidance behaviors in mice. By using
single-cell RNA-sequencing
RNA-Seq (named as an abbreviation of RNA sequencing) is a DNA sequencing, sequencing technique which uses next-generation sequencing (NGS) to reveal the presence and quantity of RNA in a biological sample at a given moment, analyzing the continu ...
, researchers have analyzed the gene expression profiles of superior colliculus neurons and identified the unique genetic markers of these circuit modules.
Other vertebrates
Optic tectum
![HE stain murine optic tectum](https://upload.wikimedia.org/wikipedia/commons/3/37/HE_stain_murine_optic_tectum.jpg)
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 ...
is the visual center in the non-mammalian brain which develops from the
alar plate
Daminozide—also known as aminozide, Alar, Kylar, SADH, B-995, B-nine, and DMASA,—is a plant growth regulator, a chemical sprayed on fruit to regulate growth, make harvest easier, and keep apples from falling off the trees before they ripen s ...
of the mesencephalon. In these other vertebrates the connections from the optic tectum are important for the recognition and reaction to various sized objects which is facilitated by excitatory optic nerve transmitters like
L-glutamate
Glutamic acid (symbol Glu or E; the ionic form is known as glutamate) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can synt ...
.
Disrupting visual experience early on in
zebrafish
The zebrafish (''Danio rerio'') is a freshwater fish belonging to the minnow family ( Cyprinidae) of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (and thus often ...
development results in a change in tectal activity. Changes in tectal activity resulted in an inability to successfully hunt and capture prey. Hypothalamus inhibitory signaling to the deep tectal
neuropil
Neuropil (or "neuropile") is any area in the nervous system composed of mostly unmyelinated axons, dendrites and glial cell processes that forms a synaptically dense region containing a relatively low number of cell bodies. The most prevalent anat ...
is important in tectal processing in zebrafish larvae. The tectal neuropil contains structures including
periventricular neuronal axons and dendrites. The neuropil also contains
GABAergic
In molecular biology and physiology, something is GABAergic or GABAnergic if it pertains to or affects the neurotransmitter GABA. For example, a synapse is GABAergic if it uses GABA as its neurotransmitter, and a GABAergic neuron produces GABA. A ...
superficial inhibitory neurons located in
stratum opticum
The retinal nerve fiber layer (RNFL) or nerve fiber layer, stratum opticum, is formed by the expansion of the fibers of the optic nerve; it is thickest near the optic disc, gradually diminishing toward the ora serrata.
As the nerve fibers pass t ...
. Instead of a large cerebral cortex, zebrafish have a relatively large optic tectum that is hypothesized to carry out some of the visual processing that the cortex performs in mammals.
Recent lesion studies have suggested that the optic tectum has no influence over higher-order motion responses like the
optomotor response
In behavioral biology, the optomotor response is an innate, orienting behavior evoked by whole-field visual motion and is common to fish and insects during locomotion, such as swimming, walking and flying. The optomotor response has algorithmic ...
or the
optokinetic response
The optokinetic response is a combination of a slow-phase and fast-phase eye movements. It is seen when an individual tracks (pursuit movement) a moving object with their eyes, which then moves out of the field of vision, a point at which their ...
, but may be more integral to lower-order cues in motion perception like in the identification of small objects.
The optic tectum is one of the fundamental components of the
vertebrate brain, existing across a range of species.
[ Maximino, 2008] Some aspects of the structure are very consistent, including a structure composed of a number of layers, with a dense input from the optic tracts to the superficial layers and another strong input conveying somatosensory input to deeper layers. Other aspects are highly variable, such as the total number of layers (from 3 in the African lungfish to 15 in the goldfish
[ Northcutt, 2002]), and the number of different types of cells (from 2 in the lungfish to 27 in the house sparrow
).
The optic tectum is closely associated with an adjoining structure called the nucleus isthmi, which has drawn a lot of interest because it evidently makes a very important contribution to tectal function.
(In the superior colliculus the like structure is termed the
parabigeminal nucleus
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 ...
). The nucleus isthmii is divided into two parts, called ''isthmus pars magnocellularis'' (Imc; "the part with the large cells") and ''isthmus pars parvocellularis'' (Ipc); "the part with the small cells"). Connections between the three areas — optic tectum, Ipc, and Imc — are topographic. Neurons in the superficial layers of the optic tectum project to corresponding points in the Ipc and Imc. The projections to the Ipc are tightly focused, while the projections to the Imc are somewhat more diffuse. Ipc gives rise to tightly focused cholinergic projections both to Imc and the optic tectum. In the optic tectum, the cholinergic inputs from Ipc ramify to give rise to terminals that extend across an entire column, from top to bottom. Imc, in contrast, gives rise to GABAergic projections to Ipc and optic tectum that spread very broadly in the lateral dimensions, encompassing most of the retinotopic map. Thus, the tectum-Ipc-Imc circuit causes tectal activity to produce recurrent feedback that involves tightly focused excitation of a small column of neighboring tectal neurons, together with global inhibition of distant tectal neurons.
The optic tectum is involved in many responses including swimming in fish, flight in birds, tongue-strikes toward prey in frogs, and fang-strikes in snakes. In some species, including fish and birds, the optic tectum, also known as the optic lobe, is one of the largest components of the brain.
In hagfish, lamprey, and shark it is a relatively small structure, but in
teleost
Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
fish it is greatly expanded, in some cases becoming the largest structure in the brain. In amphibians, reptiles, and especially birds it is also a very significant component.
In
snake
Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
s that can detect
infrared radiation
Infrared (IR), sometimes called infrared light, is electromagnetic radiation (EMR) with wavelengths longer than those of visible light. It is therefore invisible to the human eye. IR is generally understood to encompass wavelengths from around ...
, such as
python
Python may refer to:
Snakes
* Pythonidae, a family of nonvenomous snakes found in Africa, Asia, and Australia
** ''Python'' (genus), a genus of Pythonidae found in Africa and Asia
* Python (mythology), a mythical serpent
Computing
* Python (pro ...
s and
pit viper
The Crotalinae, commonly known as pit vipers,Mehrtens JM (1987). ''Living Snakes of the World in Color''. New York: Sterling Publishers. 480 pp. . crotaline snakes (from grc, κρόταλον ''krotalon'' castanet), or pit adders, are a subfa ...
s, the initial neural input is through the
trigeminal nerve
In neuroanatomy, the trigeminal nerve ( lit. ''triplet'' nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for sensation in the face and motor functions such as biting and chewin ...
instead 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 supe ...
. The rest of the processing is similar to that of the visual sense and, thus, involves the optic tectum.
Fish
Lamprey
The
lamprey
Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
has been extensively studied because it has a relatively simple brain that is thought in many respects to reflect the brain structure of early vertebrate ancestors. Inspired by the pioneering work of Carl Rovainen that began in the 1960s (see ), since the 1970s Sten Grillner and his colleagues at the Karolinska Institute in Stockholm have used the lamprey as a model system to try to work out the principles of motor control in vertebrates, starting in the spinal cord and working upward into the brain. In common with other systems (see for a historical perspective of the idea), neural circuits within the spinal cord seem capable of generating some basic rhythmic motor patterns underlying swimming, and that these circuits are influenced by specific locomotor areas in the brainstem and midbrain, that are in turn influenced by higher brain structures including the
basal ganglia
The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an extern ...
and tectum. In a study of the lamprey tectum published in 2007,
[ Saitoh et al., 2007] they found that electrical stimulation could elicit eye movements, lateral bending movements, or swimming activity, and that the type, amplitude, and direction of movement varied as a function of the location within the tectum that was stimulated. These findings were interpreted as consistent with the idea that the tectum generates goal-directed locomotion in the lamprey as shown in other species.
Birds
![SparrowTectum](https://upload.wikimedia.org/wikipedia/commons/0/09/SparrowTectum.jpg)
In birds the optic tectum is involved in flight and is one of the largest brain components. The study of
avian visual processing has enabled a greater understanding of that in mammals including humans.
See also
*
List of regions in the human brain
The human brain anatomical regions are ordered following standard neuroanatomy hierarchies. Functional, connective, and developmental regions are listed in parentheses where appropriate.
Hindbrain (rhombencephalon)
Myelencephalon
* Med ...
Additional images
file:Gray722.png, 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. (Superior colliculus visible near center.)
File:Slide5ff.JPG, Superior colliculus
File:Slide2BRA.JPG, Brainstem. Posterior view.
Notes
References
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External links
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{{DEFAULTSORT:Superior Colliculus
Midbrain
Visual system
Tectum
Corpora quadrigemina