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Perforant path | |
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Details | |
Identifiers | |
Latin | tractus perforans |
MeSH | D019580 |
NeuroNames | 2686 |
Anatomical terms of neuroanatomy |
In the brain, the perforant path or perforant pathway provides a connectional route from the entorhinal cortex [1] to all fields of the hippocampal formation, including the dentate gyrus, all CA fields (including CA1), [2] and the subiculum.
Though it arises mainly from entorhinal layers II and III, the perforant path comprises a smaller component that originates in deep layers V and VI. There is a major dichotomy with respect to the laminar origin and related terminal distribution: neurons in layer II (and possibly layer VI) project to the dentate gyrus and CA3, whereas layer III (and possibly layer V) cells project to CA1 and the subiculum via the temporoammonic pathway. [1]
In addition to playing a role in spatial memory learning generally, the temporoammonic branch (TA-CA1) of the perforant path mediates spatial memory consolidation. [3] The temporoammonic pathway has also been implicated in stress-based animal models of depression. [4]
It may also play a role in temporal lobe seizures. [5]
In rats, pyramidal and stellate cells in layer II of entorhinal cortex project through the subiculum of the hippocampus, giving rise to the name "perforant pathway". These glutamatergic fibers form a laminar pattern and terminate in the dentate gyrus and cornu ammonis 3 (CA3) region of the hippocampus. Fibers arising in the lateral portions of the entorhinal cortex show enkephalin immunoreactivity, whereas medial portions appear to contain cholecystokinin. Additionally, pyramidal cells in layer III of the entorhinal cortex send topographic projections along the perforant pathway which branch into the subiculum and CA1. [6]
In mice, the projection to CA1, and the subiculum all come primarily from EC layer III.[ citation needed ]
According to Suh et al. (2011 Science 334:1415) the projection to CA3 and dentate gyrus in mice is primarily from layer II of entorhinal cortex, and forms a trisynaptic path with hippocampus (dentate gyrus to CA3 to CA1), distinguished from the direct (monosynaptic) perforant path from Layer III of entorhinal cortex to CA1 and subiculum.[ citation needed ]
The entorhinal cortex (EC) is an area of the brain's allocortex, located in the medial temporal lobe, whose functions include being a widespread network hub for memory, navigation, and the perception of time. The EC is the main interface between the hippocampus and neocortex. The EC-hippocampus system plays an important role in declarative (autobiographical/episodic/semantic) memories and in particular spatial memories including memory formation, memory consolidation, and memory optimization in sleep. The EC is also responsible for the pre-processing (familiarity) of the input signals in the reflex nictitating membrane response of classical trace conditioning; the association of impulses from the eye and the ear occurs in the entorhinal cortex.
The hippocampus, also hippocampus proper, is a major component of the brain of humans and many other vertebrates. In the human brain the hippocampus, the dentate gyrus, and the subiculum are the components of the hippocampal formation located in the limbic system. The hippocampus plays important roles in the consolidation of information from short-term memory to long-term memory, and in spatial memory that enables navigation. In humans, and other primates the hippocampus is located in the archicortex, one of the three regions of allocortex, in each hemisphere with neural projections to the neocortex. The hippocampus, as the medial pallium, is a structure found in all vertebrates.
The dentate gyrus (DG) is one of the subfields of the hippocampus, in the hippocampal formation. The hippocampal formation is located in the temporal lobe of the brain, and includes the hippocampus subfields, and other subfields including the dentate gyrus, subiculum, and presubiculum.
In neuroanatomy, a neural pathway is the connection formed by axons that project from neurons to make synapses onto neurons in another location, to enable neurotransmission. Neurons are connected by a single axon, or by a bundle of axons known as a nerve tract, or fasciculus. Shorter neural pathways are found within grey matter in the brain, whereas longer projections, made up of myelinated axons, constitute white matter.
The entorhinal cortex (EC) is a major part of the hippocampal formation of the brain, and is reciprocally connected with the hippocampus.
Schaffer collaterals are axon collaterals given off by CA3 pyramidal cells in the hippocampus. These collaterals project to area CA1 of the hippocampus and are an integral part of memory formation and the emotional network of the Papez circuit, and of the hippocampal trisynaptic loop. It is one of the most studied synapses in the world and named after the Hungarian anatomist-neurologist Károly Schaffer.
An apical dendrite is a dendrite that emerges from the apex of a pyramidal cell. Apical dendrites are one of two primary categories of dendrites, and they distinguish the pyramidal cells from spiny stellate cells in the cortices. Pyramidal cells are found in the prefrontal cortex, the hippocampus, the entorhinal cortex, the olfactory cortex, and other areas. Dendrite arbors formed by apical dendrites are the means by which synaptic inputs into a cell are integrated. The apical dendrites in these regions contribute significantly to memory, learning, and sensory associations by modulating the excitatory and inhibitory signals received by the pyramidal cells.
The subiculum also known as the subicular complex, or subicular cortex, is the most inferior component of the hippocampal formation. It lies between the entorhinal cortex and the CA1 hippocampal subfield.
Theta waves generate the theta rhythm, a neural oscillation in the brain that underlies various aspects of cognition and behavior, including learning, memory, and spatial navigation in many animals. It can be recorded using various electrophysiological methods, such as electroencephalogram (EEG), recorded either from inside the brain or from electrodes attached to the scalp.
The stratum lucidum of the hippocampus is a layer of the hippocampus between the stratum pyramidale and the stratum radiatum. It is the tract of the mossy fiber projections, both inhibitory and excitatory from the granule cells of the dentate gyrus. One mossy fiber may make up to 37 connections to a single pyramidal cell, and innervate around 12 pyramidal cells on top of that. Any given pyramidal cell in the stratum lucidum may get input from as many as 50 granule cells.
In the hippocampus, the mossy fiber pathway consists of unmyelinated axons projecting from granule cells in the dentate gyrus that terminate on modulatory hilar mossy cells and in Cornu Ammonis area 3 (CA3), a region involved in encoding short-term memory. These axons were first described as mossy fibers by Santiago Ramón y Cajal as they displayed varicosities along their lengths that gave them a mossy appearance.
The trisynaptic circuit or trisynaptic loop is a relay of synaptic transmission in the hippocampus. The trisynaptic circuit is a neural circuit in the hippocampus, which is made up of three major cell groups: granule cells in the dentate gyrus, pyramidal neurons in CA3, and pyramidal neurons in CA1. The hippocampal relay involves three main regions within the hippocampus which are classified according to their cell type and projection fibers. The first projection of the hippocampus occurs between the entorhinal cortex (EC) and the dentate gyrus (DG). The entorhinal cortex transmits its signals from the parahippocampal gyrus to the dentate gyrus via granule cell fibers known collectively as the perforant path. The dentate gyrus then synapses on pyramidal cells in CA3 via mossy cell fibers. CA3 then fires to CA1 via Schaffer collaterals which synapse in the subiculum and are carried out through the fornix. Collectively the dentate gyrus, CA1 and CA3 of the hippocampus compose the trisynaptic loop.
Hippocampus anatomy describes the physical aspects and properties of the hippocampus, a neural structure in the medial temporal lobe of the brain. It has a distinctive, curved shape that has been likened to the sea-horse monster of Greek mythology and the ram's horns of Amun in Egyptian mythology. This general layout holds across the full range of mammalian species, from hedgehog to human, although the details vary. For example, in the rat, the two hippocampi look similar to a pair of bananas, joined at the stems. In the human and other primates, the portion of the hippocampus near the base of the temporal lobe is much broader than the part at the top. Due to the three-dimensional curvature of this structure, two-dimensional sections such as shown are commonly seen. Neuroimaging pictures can show a number of different shapes, depending on the angle and location of the cut.
The fascia dentata is the earliest stage of the hippocampal circuit. Its primary input is the perforant path from the superficial layers of entorhinal cortex. Its principal neurons are tiny granule cells which give rise to unmyelinated axons called the mossy fibers which project to the hilus and CA3. The fascia dentata of the rat contains approximately 1,000,000 granule cells. It receives feedback connections from mossy cells in the hilus at distant levels in the septal and temporal directions. The fascia dentata and the hilus together make up the dentate gyrus. As with all regions of the hippocampus, the dentate gyrus also receives GABAergic and cholinergic input from the medial septum and the diagonal band of Broca.
The name granule cell has been used for a number of different types of neurons whose only common feature is that they all have very small cell bodies. Granule cells are found within the granular layer of the cerebellum, the dentate gyrus of the hippocampus, the superficial layer of the dorsal cochlear nucleus, the olfactory bulb, and the cerebral cortex.
Sharp waves and ripples (SPW-R), also called sharp wave ripples (SWR), are oscillatory patterns produced by extremely synchronized activity of neurons in the mammalian hippocampus and neighboring regions which occur spontaneously in idle waking states or during NREM sleep. They can be observed with a variety of electrophysiological methods such as field recordings or EEG. They are composed of large amplitude sharp waves in local field potential and produced by thousands of neurons firing together within a 30–100 ms window. Within this broad time window, pyramidal cells fire only at specific times set by fast spiking GABAergic interneurons. The fast rhythm of inhibition synchronizes the firing of active pyramidal cells, each of which only fires one or two action potentials exactly between the inhibitory peaks, collectively generating the ripple pattern. SWRs have been extensively characterized by György Buzsáki and have been shown to be involved in memory consolidation in NREM sleep. Neuronal firing sequences acquired during wakefulness are replayed during SWRs.
The hippocampal subfields are four subfields CA1, CA2, CA3, and CA4 that make up the structure of the hippocampus. Regions described in the hippocampus are the head, body, and tail, and other hippocampal subfields include the dentate gyrus, the presubiculum, and the subiculum. The CA subfields use the initials of cornu ammonis, an earlier name of the hippocampus.
Early long-term potentiation (E-LTP) is the first phase of long-term potentiation (LTP), a well-studied form of synaptic plasticity, and consists of an increase in synaptic strength. LTP could be produced by repetitive stimulation of the presynaptic terminals, and it is believed to play a role in memory function in the hippocampus, amygdala and other cortical brain structures in mammals.
The supramammillary nucleus (SuM), or supramammillary area, is a thin layer of cells in the brain that lies above the mammillary bodies. It can be considered part of the hypothalamus and diencephalon. The nucleus can be divided into medial and lateral sections. The medial SuM, or SuMM, is made of smaller cells which release dopamine and give input to the lateral septal nucleus. The lateral SuM, or SuML, is made of larger cells that project to the hippocampus.
Lisa Giocomo is an American neuroscientist who is a Professor in the Department of Neurobiology at Stanford University School of Medicine. Giocomo probes the molecular and cellular mechanisms underlying cortical neural circuits involved in spatial navigation and memory.