Home LiteratureArticle Details
PMID: 12875979 Published · ppublish English Journal Article Research Support, Non-U.S. Gov't Research Support, U.S. Gov't, P.H.S.

Up-regulation of phosphorylated/activated p70 S6 kinase and its relationship to neurofibrillary pathology in Alzheimer's disease.

The American journal of pathology ·Vol. 163 ·No. 2 ·2003-08-00 ·Pages 591-607

An WL, Cowburn RF, Li L, Braak H, Alafuzoff I, Iqbal K, Iqbal IG, Winblad B, Pei JJ

Abstract

The ribosomal S6 protein kinase p70 S6 kinase is known for its role in modulating cell-cycle progression, cell size, and cell survival. In response to mitogen stimulation, p70 S6 kinase activation up-regulates ribosomal biosynthesis and enhances the translational capacity of the cell. In Alzheimer's disease (AD), there is a marked increase in total tau protein in the form of abnormally hyperphosphorylated tau (PHF-tau) in neurons with neurofibrillary tangles (NFTs). In the present study, we investigated whether p70 S6 kinase activation is associated with PHF-tau accumulation in AD. By immunohistochemistry, we found that the levels of phosphorylated p70 S6 kinase (at Thr389 or at Thr421/Ser424) were increased in accordance with the progressive sequence of neurofibrillary changes according to Braak's criteria. Confocal microscopy showed that in AD brain, phosphorylated p70 S6 kinase appeared especially in neurons that are known to later develop NFTs. This pattern of neurons showed dot-like structures of phosphorylated p70 S6 kinase and hyperphosphorylated tau, which partially correlated with rab5 (endosome marker), lamp-1 (lysosome marker), and ubiquitin (ubiquitin-proteasomal system marker). By indirect enzyme-linked immunosorbent assay, phosphorylated p70 S6 kinase (Thr389 or Thr421/Ser424), total tau, and PHF-tau were found to be significantly increased in AD brain as compared to control cases. The levels of total p70 S6 kinase and p70 S6 kinase phosphorylated at Thr421/Ser424 showed significant correlations with the levels of both total tau and PHF-tau. Regression analyses revealed a significant dependence of total tau or PHF-tau on p70 S6 kinase phosphorylated at Thr421/Ser424 rather than at Thr389. The levels of ribosomal protein S6 as well as the levels of markers for the proteolytic system were also significantly increased in AD as compared to control brain. Using a SH-SY5Y neuroblastoma cell model, we found that 100 micro mol/L zinc sulfate could induce p70 S6 kinase phosphorylation and activation, in particular at Thr421/Ser424. This up-regulation of the activated kinase resulted in an increased expression and phosphorylation of tau. Pretreatment of cells with rapamycin (an inhibitor of FRAP/mTOR which is the immediate upstream kinase of the p70 S6 kinase) attenuated the effects induced by zinc. In primary cultured neurons of rat cortical cortex, zinc sulfate treatment could repeat p70 S6 kinase phosphorylation and activation at Thr421/Ser424, followed by increased expression and phosphorylation of tau. Taken together, these data suggest that activated p70 S6 kinase could mediate an up-regulation of tau translation. The partial co-localization of phosphorylated p70 S6 kinase with rab5, lamp-1 and ubiquitin, or PHF-tau with ubiquitin suggests that the activated proteolytic system might not be sufficient to degrade the over-produced and over-phosphorylated tau protein. A p70 S6 kinase modulated up-regulation of tau translation might contribute to PHF-tau accumulation in neurons with neurofibrillary changes.

MeSH Terms
Aged Aged, 80 and over Alzheimer Disease/metabolism,pathology Animals Antibodies, Monoclonal/metabolism Biomarkers Brain/metabolism,pathology Enzyme Activation Enzyme-Linked Immunosorbent Assay Female Humans Immunohistochemistry Male Middle Aged Neurofibrils/pathology Neurons/cytology,metabolism Phosphorylation Protein Biosynthesis Ribosomal Protein S6/metabolism Ribosomal Protein S6 Kinases, 70-kDa/metabolism Signal Transduction/physiology Statistics as Topic Tumor Cells, Cultured Up-Regulation tau Proteins/genetics,metabolism
Chemicals
Antibodies, Monoclonal Biomarkers PHF-1 monoclonal antibody Ribosomal Protein S6 tau Proteins Ribosomal Protein S6 Kinases, 70-kDa
Authors & Affiliations
9 authors, click to expand affiliations / ORCID
An Wen-Lin
Division of Experimental Geriatrics, Karolinska Institutet, Neurotec, Novum, Huddinge, Sweden.
Cowburn Richard F
Li Lin
Braak Heiko
Alafuzoff Irina
Iqbal Khalid
Iqbal Inge-Grundke
Winblad Bengt
Pei Jin-Jing
References (80)
80 references, click to expand
  1. Glycogen synthase kinase-3 induces Alzheimer's disease-like phosphorylation of tau: generation of paired helical filament epitopes and neuronal localisation of the kinase.
    Neurosci Lett. 1992 Nov 23;147(1):58-62 PMID: 1336152
  2. p70s6k function is essential for G1 progression.
    Nature. 1993 May 13;363(6425):170-2 PMID: 8483501
  3. Nuclear localization of p85s6k: functional requirement for entry into S phase.
    EMBO J. 1994 Apr 1;13(7):1557-65 PMID: 8156994
  4. Cellular function of p70S6K: a role in regulating cell motility.
    Immunol Cell Biol. 2000 Aug;78(4):447-51 PMID: 10947872
  5. Application of synthetic phospho- and unphospho- peptides to identify phosphorylation sites in a subregion of the tau molecule, which is modified in Alzheimer's disease.
    J Neurosci Res. 1993 Feb 15;34(3):371-6 PMID: 8455212
  6. Enhanced neurofibrillary degeneration in transgenic mice expressing mutant tau and APP.
    Science. 2001 Aug 24;293(5534):1487-91 PMID: 11520987
  7. Increased neuronal endocytosis and protease delivery to early endosomes in sporadic Alzheimer's disease: neuropathologic evidence for a mechanism of increased beta-amyloidogenesis.
    J Neurosci. 1997 Aug 15;17(16):6142-51 PMID: 9236226
  8. Ubiquitination and abnormal phosphorylation of paired helical filaments in Alzheimer's disease.
    Mol Neurobiol. 1991;5(2-4):399-410 PMID: 1726645
  9. An array of insulin-activated, proline-directed serine/threonine protein kinases phosphorylate the p70 S6 kinase.
    J Biol Chem. 1992 Feb 15;267(5):3325-35 PMID: 1737788
  10. Mitogenesis and protein synthesis: a role for ribosomal protein S6 phosphorylation?
    Bioessays. 1994 Nov;16(11):809-15 PMID: 7840758
  11. Zinc stimulates the activity of the insulin- and nutrient-regulated protein kinase mTOR.
    Am J Physiol Endocrinol Metab. 2001 Jul;281(1):E25-34 PMID: 11404220
  12. A sequence of cytoskeleton changes related to the formation of neurofibrillary tangles and neuropil threads.
    Acta Neuropathol. 1994;87(6):554-67 PMID: 7522386
  13. Activation of the JNK/p38 pathway occurs in diseases characterized by tau protein pathology and is related to tau phosphorylation but not to apoptosis.
    J Neuropathol Exp Neurol. 2001 Dec;60(12):1190-7 PMID: 11764091
  14. Aberrant expression of mitotic cdc2/cyclin B1 kinase in degenerating neurons of Alzheimer's disease brain.
    J Neurosci. 1997 May 15;17(10):3588-98 PMID: 9133382
  15. Alzheimer's disease affects limbic nuclei of the thalamus.
    Acta Neuropathol. 1991;81(3):261-8 PMID: 1711755
  16. Phosphorylation of microtubule-associated protein tau is regulated by protein phosphatase 2A in mammalian brain. Implications for neurofibrillary degeneration in Alzheimer's disease.
    J Biol Chem. 2000 Feb 25;275(8):5535-44 PMID: 10681533
  17. Increased amount of zinc in the hippocampus and amygdala of Alzheimer's diseased brains: a proton-induced X-ray emission spectroscopic analysis of cryostat sections from autopsy material.
    J Neurosci Methods. 1997 Sep 5;76(1):53-9 PMID: 9334939
  18. Accumulation of cyclin-dependent kinase 5 (cdk5) in neurons with early stages of Alzheimer's disease neurofibrillary degeneration.
    Brain Res. 1998 Jun 29;797(2):267-77 PMID: 9666145
  19. Tangled areas of Alzheimer brain have upregulated levels of exon 10 containing tau mRNA.
    Brain Res. 1999 Jun 12;831(1-2):301-5 PMID: 10412011
  20. Ser-262 in human recombinant tau protein is a markedly more favorable site for phosphorylation by CaMKII than PKA or PhK.
    FEBS Lett. 1998 Oct 9;436(3):471-5 PMID: 9801171
  21. Identification and characterization of a mitogen-activated S6 kinase.
    Proc Natl Acad Sci U S A. 1988 Jan;85(2):406-10 PMID: 3257566
  22. Implication of brain cdc2 and MAP2 kinases in the phosphorylation of tau protein in Alzheimer's disease.
    FEBS Lett. 1992 Aug 17;308(2):218-24 PMID: 1323485
  23. Inhibition of PP-2A upregulates CaMKII in rat forebrain and induces hyperphosphorylation of tau at Ser 262/356.
    FEBS Lett. 2001 Feb 9;490(1-2):15-22 PMID: 11172803
  24. Dephosphorylation of Alzheimer paired helical filaments by protein phosphatase-2A and -2B.
    J Biol Chem. 1995 Mar 3;270(9):4854-60 PMID: 7876258
  25. Abnormal phosphorylation of the microtubule-associated protein tau (tau) in Alzheimer cytoskeletal pathology.
    Proc Natl Acad Sci U S A. 1986 Jul;83(13):4913-7 PMID: 3088567
  26. Ectopic cell cycle proteins predict the sites of neuronal cell death in Alzheimer's disease brain.
    J Neurosci. 1998 Apr 15;18(8):2801-7 PMID: 9525997
  27. Alzheimer's disease hyperphosphorylated tau sequesters normal tau into tangles of filaments and disassembles microtubules.
    Nat Med. 1996 Jul;2(7):783-7 PMID: 8673924
  28. Hydrogen peroxide activates p70(S6k) signaling pathway.
    J Biol Chem. 1999 Nov 12;274(46):32596-602 PMID: 10551813
  29. A novel pool of protein phosphatase 2A is associated with microtubules and is regulated during the cell cycle.
    J Cell Biol. 1995 Mar;128(6):1131-44 PMID: 7896877
  30. Staging of Alzheimer-related cortical destruction.
    Eur Neurol. 1993;33(6):403-8 PMID: 8307060
  31. Dynamic regulation of expression and phosphorylation of tau by fibroblast growth factor-2 in neural progenitor cells from adult rat hippocampus.
    J Neurosci. 1999 Jul 1;19(13):5245-54 PMID: 10377336
  32. Extracellular zinc activates p70 S6 kinase through the phosphatidylinositol 3-kinase signaling pathway.
    J Biol Chem. 2000 Aug 25;275(34):25979-84 PMID: 10851233
  33. Dissection of the protein kinase cascade by which nerve growth factor activates MAP kinases.
    Nature. 1991 Sep 12;353(6340):170-3 PMID: 1716348
  34. A single gene encodes two isoforms of the p70 S6 kinase: activation upon mitogenic stimulation.
    Proc Natl Acad Sci U S A. 1992 May 1;89(9):4052-6 PMID: 1570332
  35. Localization of active forms of C-jun kinase (JNK) and p38 kinase in Alzheimer's disease brains at different stages of neurofibrillary degeneration.
    J Alzheimers Dis. 2001 Feb;3(1):41-48 PMID: 12214071
  36. Monoclonal antibody PHF-1 recognizes tau protein phosphorylated at serine residues 396 and 404.
    J Neurosci Res. 1994 Dec 15;39(6):669-73 PMID: 7534834
  37. Phosphorylation of tau at both Thr 231 and Ser 262 is required for maximal inhibition of its binding to microtubules.
    Arch Biochem Biophys. 1998 Sep 15;357(2):299-309 PMID: 9735171
  38. Neuropathological stageing of Alzheimer-related changes.
    Acta Neuropathol. 1991;82(4):239-59 PMID: 1759558
  39. Chloroquine myopathy suggests that tau is degraded in lysosomes: implication for the formation of paired helical filaments in Alzheimer's disease.
    Neurosci Res. 1998 May;31(1):1-8 PMID: 9704973
  40. Silver impregnation of Alzheimer's neurofibrillary changes counterstained for basophilic material and lipofuscin pigment.
    Stain Technol. 1988 Jul;63(4):197-200 PMID: 2464205
  41. Cell cycle markers in the hippocampus in Alzheimer's disease.
    Acta Neuropathol. 1997 Jul;94(1):6-15 PMID: 9224524
  42. Distribution of active glycogen synthase kinase 3beta (GSK-3beta) in brains staged for Alzheimer disease neurofibrillary changes.
    J Neuropathol Exp Neurol. 1999 Sep;58(9):1010-9 PMID: 10499443
  43. Identification and localization of a tau peptide to paired helical filaments of Alzheimer disease.
    Proc Natl Acad Sci U S A. 1989 Jul;86(14):5646-50 PMID: 2501795
  44. Microtubule-associated protein tau. Abnormal phosphorylation of a non-paired helical filament pool in Alzheimer disease.
    J Biol Chem. 1993 Nov 15;268(32):24374-84 PMID: 8226987
  45. TGF-beta inhibits p70 S6 kinase via protein phosphatase 2A to induce G(1) arrest.
    Genes Dev. 2000 Dec 15;14(24):3093-101 PMID: 11124802
  46. Constitutive Cdc25B tyrosine phosphatase activity in adult brain neurons with M phase-type alterations in Alzheimer's disease.
    Neuroscience. 2001;105(3):639-50 PMID: 11516829
  47. Selective phosphorylation of adult tau isoforms in mature hippocampal neurons exposed to fibrillar A beta.
    Mol Cell Neurosci. 1997;9(3):220-34 PMID: 9245504
  48. Role of protein phosphatase-2A and -1 in the regulation of GSK-3, cdk5 and cdc2 and the phosphorylation of tau in rat forebrain.
    FEBS Lett. 2000 Nov 17;485(1):87-93 PMID: 11086171
  49. Signals of oxidant-induced cardiomyocyte hypertrophy: key activation of p70 S6 kinase-1 and phosphoinositide 3-kinase.
    J Pharmacol Exp Ther. 2002 Mar;300(3):1101-10 PMID: 11861821
  50. Phosphatase activity toward abnormally phosphorylated tau: decrease in Alzheimer disease brain.
    J Neurochem. 1995 Aug;65(2):732-8 PMID: 7616230
  51. Up-regulation of cell division cycle (cdc) 2 kinase in neurons with early stage Alzheimer's disease neurofibrillary degeneration.
    Acta Neuropathol. 2002 Oct;104(4):369-76 PMID: 12200623
  52. Substrate recognition determinants of the mitogen-activated 70K S6 kinase from rat liver.
    J Biol Chem. 1992 Feb 15;267(5):3074-8 PMID: 1737763
  53. Defective brain microtubule assembly in Alzheimer's disease.
    Lancet. 1986 Aug 23;2(8504):421-6 PMID: 2874414
  54. Identification of kinase-phosphatase signaling modules composed of p70 S6 kinase-protein phosphatase 2A (PP2A) and p21-activated kinase-PP2A.
    J Biol Chem. 1999 Jan 8;274(2):687-92 PMID: 9873003
  55. Up-regulation of mitogen-activated protein kinases ERK1/2 and MEK1/2 is associated with the progression of neurofibrillary degeneration in Alzheimer's disease.
    Brain Res Mol Brain Res. 2002 Dec 30;109(1-2):45-55 PMID: 12531514
  56. Levels of normal and abnormally phosphorylated tau in different cellular and regional compartments of Alzheimer disease and control brains.
    FEBS Lett. 1994 Aug 29;351(1):80-4 PMID: 8076698
  57. Cloning and expression of two human p70 S6 kinase polypeptides differing only at their amino termini.
    Mol Cell Biol. 1991 Nov;11(11):5541-50 PMID: 1922062
  58. Silver staining of Alzheimer's neurofibrillary changes by means of physical development.
    Acta Morphol Acad Sci Hung. 1971;19(1):1-8 PMID: 4107507
  59. Role of abnormally phosphorylated tau in the breakdown of microtubules in Alzheimer disease.
    Proc Natl Acad Sci U S A. 1994 Jun 7;91(12):5562-6 PMID: 8202528
  60. Early phosphorylation of tau in Alzheimer's disease occurs at Ser-202 and is preferentially located within neurites.
    Neuroreport. 1994 Nov 21;5(17):2358-62 PMID: 7533559
  61. Using structure to define the function of phosphoinositide 3-kinase family members.
    FEBS Lett. 1997 Jun 23;410(1):91-5 PMID: 9247130
  62. Brain ubiquitin is markedly elevated in Alzheimer disease.
    Brain Res. 1991 Dec 6;566(1-2):146-51 PMID: 1814531
  63. Distribution, levels, and activity of glycogen synthase kinase-3 in the Alzheimer disease brain.
    J Neuropathol Exp Neurol. 1997 Jan;56(1):70-8 PMID: 8990130
  64. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding.
    Anal Biochem. 1976 May 7;72:248-54 PMID: 942051
  65. Phosphorylation and activation of p70s6k by PDK1.
    Science. 1998 Jan 30;279(5351):707-10 PMID: 9445476
  66. Amyloid-beta injection in rat amygdala alters tau protein but not mRNA expression.
    Exp Neurol. 2000 Mar;162(1):158-70 PMID: 10716896
  67. Microtubule-associated protein tau. A component of Alzheimer paired helical filaments.
    J Biol Chem. 1986 May 5;261(13):6084-9 PMID: 3084478
  68. Ribosomal S6 kinase signaling and the control of translation.
    Exp Cell Res. 1999 Nov 25;253(1):100-9 PMID: 10579915
  69. Epidemiology of Alzheimer's disease. Issues of etiology and validity.
    Acta Neurol Scand Suppl. 1993;145:1-70 PMID: 8333250
  70. Defective ubiquitination of cerebral proteins in Alzheimer's disease.
    J Neurosci Res. 2000 Oct 15;62(2):302-10 PMID: 11020223
  71. Imbalances of trace elements related to oxidative damage in Alzheimer's disease brain.
    Neurotoxicology. 1998 Jun;19(3):339-45 PMID: 9621340
  72. Formation of neurofibrillary tangles in P301l tau transgenic mice induced by Abeta 42 fibrils.
    Science. 2001 Aug 24;293(5534):1491-5 PMID: 11520988
  73. In situ hybridization of calcium/calmodulin dependent protein kinase II and tau mRNAs; species differences and relative preservation in Alzheimer's disease.
    Brain Res Mol Brain Res. 1992 Jan;12(1-3):85-94 PMID: 1312209
  74. Cdc2 phosphorylation of nucleolin demarcates mitotic stages and Alzheimer's disease pathology.
    Neurobiol Aging. 2001 Jul-Aug;22(4):517-28 PMID: 11445251
  75. Copper, iron, and zinc imbalances in severely degenerated brain regions in Alzheimer's disease: possible relation to oxidative stress.
    J Neurol Sci. 1996 Nov;143(1-2):137-42 PMID: 8981312
  76. A serine/threonine proline kinase activity is included in the tau protein kinase fraction forming a paired helical filament epitope.
    Neurosci Lett. 1991 Jul 22;128(2):195-8 PMID: 1658692
  77. Accumulation of tau in autophagic vacuoles in chloroquine myopathy.
    J Neuropathol Exp Neurol. 1998 Jul;57(7):664-73 PMID: 9690670
  78. Occurrence of neuropil threads in the senile human brain and in Alzheimer's disease: a third location of paired helical filaments outside of neurofibrillary tangles and neuritic plaques.
    Neurosci Lett. 1986 Apr 24;65(3):351-5 PMID: 2423928
  79. Hyperphosphorylation and accumulation of neurofilament proteins in Alzheimer disease brain and in okadaic acid-treated SY5Y cells.
    FEBS Lett. 2001 Oct 19;507(1):81-7 PMID: 11682063
  80. Brain levels of microtubule-associated protein tau are elevated in Alzheimer's disease: a radioimmuno-slot-blot assay for nanograms of the protein.
    J Neurochem. 1992 Aug;59(2):750-3 PMID: 1629745
Article Info
Journal
The American journal of pathology
Abbr.
Am J Pathol
ISSN
0002-9440
Published
2003-08-00
Pages
591-607
Language
English
Region
United States
NLM ID
0370502
PMCID
PMC1868198
Subset
IM
Grants
NIA NIH HHS · R01 AG019158 · United States
NIA NIH HHS · AG08076 · United States
NIA NIH HHS · AG19158 · United States
Corrections
ErratumIn
-
Analysis Services
Analysis Services

Contact

No. 2 Wenbo Road, Zhangqiu District, Jinan, Shandong

Qilu Normal University · Genelibs Bioinformatics Lab

750 Shunhua Rd, Jinan

2F, Bldg F, University Science Park

Tel: 0531-88819269

WeChat Official Account

Follow our WeChat subscription account for real-time updates and the latest in medical and biological research.


Business Email

E-mail: product@genelibs.com