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|Astrocytes can be visualized in culture because, like other glia, they express glial fibrillary acidic protein.|
Astrocytes (also known collectively as astroglia) are characteristic star-shaped glial cells in the brain and spinal cord. They perform many functions, including biochemical support of endothelial cells which form the blood-brain barrier, provision of nutrients to the nervous tissue, maintenance of extracellular ion balance, and a principal role in the repair and scarring process of the brain and spinal cord following traumatic injuries.
Research since the mid-1990s has shown that astrocytes propagate intercellular Ca2+ waves over long distances in response to stimulation, and, similar to neurons, release transmitters (called gliotransmitters) in a Ca2+-dependent manner. Data suggest that astrocytes also signal to neurons through Ca2+-dependent release of glutamate. Such discoveries have turned astrocyte research into a rapidly growing field of neuroscience.
DescriptionEditAstrocytes are a sub-type of glial cells in the central nervous system. They are also known as astrocytic glial cells. Star-shaped, their many processes envelope synapses made by neurons. Astrocytes are classically identified using histological analysis; many of these cells express the intermediate filament glial fibrillary acidic protein (GFAP). Three forms of astrocytes exist in the CNS, fibrous, protoplasmic and radial. The fibrous glia are usually located within white matter, have relatively few organelles, and exhibit long unbranched cellular processes. This type often has "vascular feet" that physically connect the cells to the outside of capillary wall when they are in close proximity to them. The protoplasmic glia are found in grey matter tissue, possess a larger quantity of organelles, and exhibit short and highly branched cellular processes. Lastly, the radial glia are disposed in a plane perpendicular to axis of ventricles. One of their processes about the pia mater, while the other is deeply buried in gray matter. Radial glia is mostly present during development, playing a role in neuron migration. Mueller cells of retina and Bergmann glia cells of cerebellar cortex represent an exception, being present still during adulthood. When in proximity to the pia mater, all three forms of astrocytes send out process to form the pia-glial membrane.
Previously in medical science, the neuronal network was considered the only important one, and astrocytes were looked upon as gap fillers. More recently, the function of astrocytes has been reconsidered, and are now thought to play a number of active roles in the brain, including the secretion or absorption of neural transmitters and maintenance of the blood-brain barrier. Following on this idea the concept of a "tripartite synapse" has been proposed, referring to the tight relationship occurring at synapses among a presynaptic element, a postsynaptic element and a glial element..
- Structural: involved in the physical structuring of the brain.
- Metabolic support: they provide neurons with nutrients such as lactate.
- Blood-brain barrier: the astrocyte end-feet encircling endothelial cells were thought to aid in the maintenance of the blood-brain barrier, but recent research indicates that they do not play a substantial role; instead it is the tight junctions and basal lamina of the cerebral endothelial cells that play the most substantial role in maintaining the barrier. However, it has recently been shown that astrocyte activity is linked to blood flow in the brain, and that this is what is actually being measured in fMRI.
- Transmitter reuptake and release: astrocytes express plasma membrane transporters such as glutamate transporters for several neurotransmitters, including glutamate, ATP and GABA. More recently, astrocytes were shown to release glutamate or ATP in a vesicular, Ca2+-dependent manner.
- Regulation of ion concentration in the extracellular space: astrocytes express potassium channels at a high density. When neurons are active, they release potassium, increasing the local extracellular concentration. Because astrocytes are highly permeable to potassium, they rapidly clear the excess accumulation in the extracellular space. If this function is interfered with, the extracellular concentration of potassium will rise, leading to neuronal depolarization by the Goldman equation. Abnormal accumulation of extracellular potassium is well known to result in epileptic neuronal activity.
- Modulation of synaptic transmission: in the supraoptic nucleus of the hypothalamus, rapid changes in astrocyte morphology have been shown to affect heterosynaptic transmission between neurons. In the hippocampus, astrocytes suppress synaptic transmission by releasing ATP, which is hydrolyzed by ectonucliotidases to yield adenosine. Adenosine acts on neuronal adenosine receptors to inhibit synaptic transmission, thereby increasing the dynamic range available for LTP.
- Vasomodulation: astrocytes may serve as intermediaries in neuronal regulation of blood flow.
- Promotion of the myelinating activity of oligodendrocytes: electrical activity in neurons causes them to release ATP, which serves as an important stimulus for myelin to form. Surprisingly, the ATP does not act directly on oligodendrocytes. Instead it causes astrocytes to secrete cytokine leukemia inhibitory factor (LIF), a regulatory protein that promotes the myelinating activity of oligodendrocytes. This suggest that astrocytes have an executive-coordinating role in the brain.
- Nervous system repair: upon injury to nerve cells within the central nervous system, astrocytes become phagocytic to ingest the injured nerve cells. The astrocytes then fill up the space to form a glial scar, repairing the area and replacing the CNS cells that cannot regenerate.
Recent studies have shown that astrocytes play an important function in the regulation of neural stem cells. Research from the Schepens Eye Research Institute at Harvard shows the human brain to abound in neural stem cells, which are kept in a dormant state by chemical signals (ephrin-A2 and ephrin-A3) from the astrocytes. The astrocytes are able to activate the stem cells to transform into working neurons by dampening the release of ephrin-A2 and ephrin-A3.
Furthermore, studies are underway to determine whether astroglia play an instrumental role in depression, based on the link between diabetes and depression. Altered CNS glucose metabolism is seen in both these conditions, and the astroglial cells are the only cells with insulin receptors in the brain.
An increase in intracellular calcium concentration can propagate outwards through this syncytium. Mechanisms of calcium wave propagation include diffusion of IP3 through gap junctions and extracellular ATP signalling. Calcium elevations are the primary known axis of activation in astrocytes, and are necessary and sufficient for some types of astrocytic glutamate release.
There are several different ways to classify astrocytes:
by Lineage and antigenic phenotypeEdit
These have been established by classic work by Raff et al. in early 1980s on Rat optic nerves.
- Type 1: Antigenically Ran2+, GFAP+, FGFR3+, A2B5- thus resembling the "type 1 astrocyte" of the postnatal day 7 rat optic nerve. These can arise from the tripotential glial restricted precursor cells (GRP), but not from the bipotential O2A/OPC (oligodendrocyte, type 2 astrocyte precursor, also called Oligodendrocyte progenitor cell) cells.
- Type 2: Antigenically A2B5+, GFAP+, FGFR3-, Ran 2-. These cells can develop in vitro from the either tripotential GRP (probably via O2A stage) or from bipotential O2A cells (which some people[attribution needed] think may in turn have been derived from the GRP) or in vivo when the these progenitor cells are transplanted into lesion sites (but probably not in normal development, at least not in the rat optic nerve). Type-2 astrocytes are the major astrocytic component in postnatal optic nerve cultures that are generated by O2A cells grown in the presence of fetal calf serum but are not thought to exist in vivo (Fulton et al., 1992).
by Anatomical ClassificationEdit
- Protoplasmic: found in grey matter and have many branching processes whose end-feet envelop synapses. Some protoplasmic astrocytes are generated by multipotent subventricular zone progenitor cells.
- Gömöri-positive astrocytes. These are a subset of protoplasmic astrocytes that contain numerous cytoplasmic inclusions, or granules, that stain positively with Gömöri's chrome-alum hematoxylin stain. It is now known that these granules are formed from the remnants of degenerating mitochondria engulfed within lysosomes , Some type of oxidative stress appears to be responsible for the mitochondrial damage within these specialized astrocytes. Gömöri-positive astrocytes are much more abundant within the arcuate nucleus of the hypothalamus and in the hippocampus than in other brain regions. They may have a role in regulating the response of the hypothalamus to glucose .
- Fibrous: found in white matter and have long thin unbranched processes whose end-feet envelop nodes of Ranvier. Some fibrous astrocytes are generated by radial glia.
by Transporter/receptor classificationEdit
- GluT type: express glutamate transporters (EAAT1/SLC1A3 and EAAT2/SLC1A2) and respond to synaptic release of glutamate by transporter currents
- GluR type: express glutamate receptors (mostly mGluR and AMPA type) and respond to synaptic release of glutamate by channel-mediated currents and IP3-dependent Ca2+ transients
Bergmann glia, a type of glia also known as radial epithelial cells (as named by Camillo Golgi) or Golgi epithelial cells (GCEs; not to be mixed up with Golgi cells), are astrocytes in the cerebellum that have their cell bodies in the Purkinje cell layer and processes that extend into the molecular layer, terminating with bulbous endfeet at the pial surface. Bergmann glia express high densities of glutamate transporters that limit diffusion of the neurotransmitter glutamate during its release from synaptic terminals. Besides their role in early development of the cerebellum, Bergmann glia are also required for the pruning or addition of synapses.
- ↑ Fiacco TA, Agulhon C, McCarthy KD (October 2008). Sorting out Astrocyte Physiology from Pharmacology. Annu. Rev. Pharmacol. Toxicol. 49: 151.
- ↑ The Brain From Top To Bottom. Thebrain.mcgill.ca. URL accessed on 2008-11-29.
- ↑ Kolb & Whishaw: Fundamentals of Human Neuropsychology, 2008
- ↑ Araque A, Parpura V, Sanzgiri RP, Haydon PG (1999). Tripartite synapses: glia, the unacknowledged partner.. Trends in Neuroscience 22: 208–215.
- ↑ Swaminathan N (2008). Brain-scan mystery solved. Scientific American Mind Oct-Nov.
- ↑ Santello M, Volterra A (2008). Synaptic modulation by astrocytes via Ca(2+)-dependent glutamate release.. Neuroscience Mar 22 (1): 253.
- ↑ Piet R, Vargová L, Syková E, Poulain D, Oliet S (2004). Physiological contribution of the astrocytic environment of neurons to intersynaptic crosstalk. Proc Natl Acad Sci USA 101 (7): 2151–5.
- ↑ Pascual O, Casper KB, Kubera C, Zhang J, Revilla-Sanchez R, Sul JY, Takano H, Moss SJ, McCarthy K, Haydon PG (2005). Astrocytic purinergic signaling coordinates synaptic networks. Science 310 (5745): 113–6.
- ↑ Parri R, Crunelli V (2003). An astrocyte bridge from synapse to blood flow. Nat Neurosci 6 (1): 5–6.
- ↑ Ishibashi T, Dakin K, Stevens B, Lee P, Kozlov S, Stewart C, Fields R (2006). Astrocytes promote myelination in response to electrical impulses. Neuron 49 (6): 823–32.
- ↑ Bennett M, Contreras J, Bukauskas F, Sáez J (2003). New roles for astrocytes: gap junction hemichannels have something to communicate. Trends Neurosci 26 (11): 610–7.
- ↑ Newman, EA.(2001)"Propagation of intercellular calcium waves in retinal astrocytes and Müller cells."J Neurosci. 21(7):2215-23
- ↑ Parpura V, Haydon P (2000). Physiological astrocytic calcium levels stimulate glutamate release to modulate adjacent neurons. Proc Natl Acad Sci USA 97 (15): 8629–34.
- ↑ Levison SW, Goldman JE (February 1993). Both oligodendrocytes and astrocytes develop from progenitors in the subventricular zone of postnatal rat forebrain. Neuron 10 (2): 201–12.
- ↑ Zerlin M, Levison SW, Goldman JE (November 1995). Early patterns of migration, morphogenesis, and intermediate filament expression of subventricular zone cells in the postnatal rat forebrain. J. Neurosci. 15 (11): 7238–49.
- ↑ Brawer,JR. (1994) "Composition of Gomori-positive inclusions in astrocytes of the hypothalamic arcuate nucleus" Anatomical Record 240: 407-415. PMID 7825737
- ↑ Young JK, McKenzie JC (2004) "GLUT2 immunoreactivity in Gömöri-positive astrocytes of the hypothalamus."J. Histochemistry & Cytochemistry 52: 1519-1524 PMID
- ↑ Marty, N. (2005) "Regulation of glucagon secretion by glucose transporter type 2 (glut2) and astrocyte-dependent glucose sensors" J. Clinical Investigation 115: 3545.
- ↑ Memorial University of Newfoundland - Anatomy at MUN nerve/neuron
- ↑ Choi BH, Lapham LW (June 1978). Radial glia in the human fetal cerebrum: a combined Golgi, immunofluorescent and electron microscopic study. Brain Res. 148 (2): 295–311.
- ↑ Schmechel DE, Rakic P (June 1979). A Golgi study of radial glial cells in developing monkey telencephalon: morphogenesis and transformation into astrocytes. Anat. Embryol. 156 (2): 115–52.
- ↑ Misson JP, Edwards MA, Yamamoto M, Caviness VS (November 1988). Identification of radial glial cells within the developing murine central nervous system: studies based upon a new immunohistochemical marker. Brain Res. Dev. Brain Res. 44 (1): 95–108.
- ↑ Voigt T (November 1989). Development of glial cells in the cerebral wall of ferrets: direct tracing of their transformation from radial glia into astrocytes. J. Comp. Neurol. 289 (1): 74–88.
- ↑ Goldman SA, Zukhar A, Barami K, Mikawa T, Niedzwiecki D (August 1996). Ependymal/subependymal zone cells of postnatal and adult songbird brain generate both neurons and nonneuronal siblings in vitro and in vivo. J. Neurobiol. 30 (4): 505–20.
- ↑ Riquelme R, Miralles C, De Blas A (15 December 2002). Bergmann glia GABA(A) receptors concentrate on the glial processes that wrap inhibitory synapses. J. Neurosci. 22 (24): 10720–30.
- ↑ Yamada K, Watanabe M (2002). Cytodifferentiation of Bergmann glia and its relationship with Purkinje cells. Anatomical science international / Japanese Association of Anatomists 77 (2): 94–108.
- "Astrocytes" at Society for Neuroscience
- The Department of Neuroscience at Wikiversity
- NIF Search - Astrocyte via the Neuroscience Information Framework
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