Literature: A-K ; L-Z ; subject area
Compiled by: Julian Thorpe
Neurofilaments
|
Please Note: Due to time constraints, the text part of this page has not been updated for some time. However, references are added reasonably frequently. |
glial filaments ; peripherin ; internexin ; references
General
Neurofilaments (NFs) are the (Type IV) intermediate filaments found in neurons.
 | N.B. See Steinert (1996) and Julien and Mushynski (1998) for reviews on 'Intermediate Filaments in Health and Disease'. |
| Most abundant structures in most neurons. |
| Work has suggested a critical role for neurofilaments in the development
and maintenance of axonal calibre, crucial for the conduction velocity
of axons (e.g. Cleveland et al., 1991; Yamasaki et al., 1992
and Eyer and Peterson, 1994). |
| They are formed from a triplet of three different-sized protein subunits termed NF-L , NF-M and NF-H (abbreviations for light [68kD], medium [160kD] and heavy [200kD]). |
| NF-M and NF-H have extended C-terminal domains and are thus unusually large for intermediate filaments. |
| Occur as lengthier filaments in axons compared with dendrites [in cultured hippocampal neurons: Benson et al., 1996). |
| The C-terminal domains are glutamic acid-enriched and can be phosphorylated on many residues (see refs below). They protrude sideways to connect with other filaments, microtubules or vesicles. |
| The C-terminal region of NF-H contains a unique functional domain
consisting of 43 repeat Lys-Ser-Pro (KSP) motifs with either 3 or 5 amino
acid spacers between them (see NF-H sequence data
). |
| NF-L and NF-Mfirst expressed shortly after initiation of axonal outgrowth with expression of NF-Hoccurring a few days later (in cultured hippocampal neurons: Benson et al., 1996 ). |
| In cultured rat brain neurons NF proteins were found to disappear and then NF-L and NF-M reappeared first, followed by dephosphorylated, and then phosphorylated, NF-H. The appearance of phosphorylated NF-H coincided with synapse formation (at around 4 days; Tokuoka et al., 2000 ). The NF-H phosphorylation was shown to be stimulated by brain-derived neurotrophic factor (BDNF) and neurotrophin 3 (NT3). |
| The C-terminal domains are glutamic acid-enriched and can be phosphorylated on many residues (see refs below) by a large number of protein kinases . |
| Phosphorylation of NFs probably has a role in the establishment of neurofilament density and axonal calibre. There is also growing evidence (based on transgenic mouse studies ) that neurofilaments can affect the dynamics and perhaps the function of other cytoskeletal elements, such as microtubules and actin filaments . Perturbations in phosphorylation or in metabolism of neurofilaments are frequently observed in neurodegenerative diseases. |
| Most of the neurofilament phosphorylation sites, located in tail regions of NF-M and NF-H, consist of the repeat sequence motif, Lys-Ser-Pro (KSP). |
| The highly-phosphorylated Lys-Ser-Pro (KSP) region of NF-H is variable between species, while other regions are highly homologous ( Breen et al., 1989). Figlewicz et al. (1993) have identified an allelic variant of human NF-H containing an additional KSP motif. |
| Lee et al. (1988a and 1988b) have shown that there is a conserved major NF phosphorylation site within mammalian (and invertebrate) species. An antigenic determinant of this region was shown to exist in neurofibrillary tangles and in the MAP proteins MAP2 and tau . |
| It has been proposed that accumulations of variably-phosphorylated NFs represent the earliest cytoskeletal alteration associated with dystrophic neurite formation in AD (Dickson et al., 1999). These authors suggested that dystrophic neurites might 'mature' through NF-abundant forms through to the neurites containing the profoundly altered tau -immunoreactive filaments and that this was caused by amyloid plaques exerting a physically-damaging effect upon neighbouring axons. |
| NF-H is usually hypophosphorylated in the neuronal perikaryon but extensively hyperphosphorylated in axons. |
| Aberrant hyperphosphorylation of perikaryal NF-H is a common feature of many neurological diseases. Giasson et al. (1997) have shown that activation of stress-activated protein kinases (SAPKs ) by an upstream activator (MEKK-1) caused extensive NFH phosphorylation. SAPK was localised in the neurites as well as the cell body and they suggested that it could be involved in the phosphorylation of NFH in neurites (neuritic NFH is highly-phosphorylated despite a demonstrated absence of cyclin-dependent kinase 5 activity in these neurons). |
| Increased levels of phosphorylated NF-H in AD brain: Hashimoto et al. (1999) have shown that the relative amount of phosphorylated NF-H was increased in AD brains compared with controls (there were no apparent changes in NF-L and NF-M). |
| Anti-neurofilament serum antibodies: The work of Oron et al. (1997) on a rat animal model system suggests a role for serum antibodies (against phosphorylated epitopes highly enriched in the heavy neurofilament protein NF-H) in the neurodegeneration of cholinergic neurons in AD. |
| Jung et al. (2000) demonstrated that phosphorylation of the C-terminus of NF-H correlated with decreased neurofilament axonal transport velocity and that this phosphorylation therefore mediated the regulation of axonal transport. |
| Brownlees et al. (2000) showed that stress activated protein kinase 1b (SAPK1b) phosphorylates (multiple phosphorylation sites on) NF-H. They also showed that glutamate treatment induced activation of the SAPK1b and increased NF-H phosphorylation in the (primary cortical neuronal) cell bodies. |
| Ahlijanian et al. (2000) used transgenic mice (overexpressing human p25, which activates cdK5 ) to show hyperphosphorylation of tau and neurofilaments by cdk5. This latter was accompanied by cytoskeletal disruption. |
| Nakamura et al. (2000) demonstrated that bovine NF-L was phosphorylated by PKA at Ser41, Ser55, and Ser62 in the head region. They also showed that Ser55 phosphorylation could be modulated by okadaic acid-sensitive phosphatases . |
| Tokuoka et al. (2000) showed that NF-H phosphorylation (in cultured rat brain neurons) was stimulated by brain-derived neurotrophic factor (BDNF) and neurotrophin 3 (NT3) and correlated with synapse formation. The BDNF-stimulated phosphorylation (and thus synapse formation) was mediated, at least in part, through activation of cdk5 . |
| Cdk5 and ERK1/2 are associated with complexes of NF proteins (and tubulins and tau ; Veeranna et al., 2000). |
| Strong et al. (1994) used rabbits to test the effects of aluminium neurotoxicity and found that NF-L and NF-M messenger RNA levels were reduced in direct proportion to the extent of NF inclusion formation. |
| Tu et al. (1997) found that transgenic mice expressing a fusion protein (composed of NF-H fused to beta-galactosidase) developed inclusions in neurons throughout the CNS. These contained massive filamentous aggregates of the endogenous NF proteins and the fusion protein which resembled the NF-rich Lewy bodies of Parkinson's disease and Lewy Body dementia. In hippocampal neurons they found that the inclusions which entrapped organelles (type II) led to neuronal cell death. |
| Trojanowski et al. (1998a) have reviewed abnormal protein aggregations in Parkinson's disease and Lewy body dementia. They state that Lewy bodies occur predominantly in the neocortical neurons in a subtype of AD known as the LB variant of AD (LBVAD) and that "aggregated neurofilament subunits and alpha-synuclein are major protein components of LBs, and these inclusions may contribute mechanistically to the degeneration of neurons in PD, DLB and LBVAD." |
| Abnormal NF accumulations are a prominent feature of brain trauma.These accumulations are predominantly dephosphorylated (in swine: Chen et al., 1999). |
| It has been proposed that accumulations of variably-phosphorylated NFs represent the earliest cytoskeletal alteration associated with dystrophic neurite formation in AD (Dickson et al., 1999). These authors suggested that dystrophic neurites might 'mature' through NF-abundant forms through to the neurites containing the profoundly altered tau -immunoreactive filaments and that this was caused by amyloid plaques exerting a physically-damaging effect upon neighbouring axons. |
| Norlund et al. (1999) showed that there were increased levels of transglutaminase-induced epsilon-(gamma-glutamyl)lysine bonds (in AD brain) in the NF-M and NF-H proteins (and in tau , which contributes towards the stabilisation of NFT s and their resistance to degradation and proteolytic digestion). |
| Increased titres of anti-NF antibodies have been reported in neurodegenerative disorders and it has been suggested that they might be pathogenic. |
| Terry Lorenzo et al. (2000) showed that NF-L (purified from bovine brain cortex plasma membranes) was a major protein phosphatase 1 (PP1)-binding target which also inhibited PP1, but not PP2A, activity. They suggested that NF-L might target PP1 in membranes and the cytoskeleton of mammalian neurons. |
| Ray et al. (2000) have used PC12 cells to show that the upregulation of calpain (a Ca2+-dependent cysteine protease) by oxidative stress and calcium influx, resulted in NF-L degradation (and apoptosis). |
| Sjögren et al. (2000) investigated the CSF levels of
(tau
and) NF-L
in patients with frontotemporal dementia (FTD) and other common
dementia disorders (including early- and late-onset AD).Their results
suggested an involvement of NF-L primarily in FTD and tau in early-onset
AD. They suggested that their observed increases in CSF NF-L found in late-onset
AD might reflect white-matter degeneration. |
| These are the specific intermediate filaments of the glial cells (latter are found in mammalian brain and appear to support neuronal cells). The glial filaments are composed of glial fibrillary acidic protein (GFAP). |
| Terryberry et al. (1998) found that some (33%) of their AD patients studied had glial fibrillary acidic protein autoantibodies (in sera and cerebrospinal fluid). |
| Is a type III intermediate filament found in peripheral and certain CNS neurons (contains a phosphotyrosine residue: Angelastro et al., 1998). |
| Found to co-immunoprecipitate with (alpha) internexin (in cultured
rat neurons): Athlan et al., 1997. |
| Is present in all neurons at every developmental stage (as long filaments in axons and short fragments in dendrites of cultured hippocampal neurons: Benson et al., 1996 ). |
| May serve as a link between cytoskeletal elements in dendritic shafts and spines (Benson et al., 1996). |
| Co-immunoprecipitates with (alpha) internexin (in cultured rat neurons): Athlan et al., 1997). |
Some Related References
(for those referring more specifically to tau, see the
neurofibrillary tangle
page)
(These journals include: Neurology, Neurobiology of Disease, Journal of Neurochemistry, Alzheimer's Disease Review)
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