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Volumn 21, Issue 8, 2012, Pages 1824-1834

Defective glucocorticoid hormone receptor signaling leads to increased stress and anxiety in a mouse model of Angelman syndrome

Author keywords

[No Author keywords available]

Indexed keywords

CORTICOSTERONE; GLUCOCORTICOID RECEPTOR; PARVALBUMIN; UBIQUITIN PROTEIN LIGASE E3; UBIQUITIN PROTEIN LIGASE E3A; UNCLASSIFIED DRUG;

EID: 84859256771     PISSN: 09646906     EISSN: 14602083     Source Type: Journal    
DOI: 10.1093/hmg/ddr614     Document Type: Article
Times cited : (51)

References (54)
  • 1
    • 0037328861 scopus 로고    scopus 로고
    • Angelman syndrome: a review of the clinical and genetic aspects
    • Clayton-Smith, J. and Laan, L. (2003) Angelman syndrome: a review of the clinical and genetic aspects. J. Med. Genet., 40, 87-95.
    • (2003) J. Med. Genet. , vol.40 , pp. 87-95
    • Clayton-Smith, J.1    Laan, L.2
  • 2
    • 12744269033 scopus 로고    scopus 로고
    • Neurological aspects of the Angelman syndrome
    • Williams, C.A. (2005) Neurological aspects of the Angelman syndrome. Brain Dev., 27, 88-94.
    • (2005) Brain Dev. , vol.27 , pp. 88-94
    • Williams, C.A.1
  • 5
    • 0031012849 scopus 로고    scopus 로고
    • UBE3A/E6-AP mutations cause Angelman syndrome
    • Kishino, T., Lalande, M. and Wagstaff, J. (1997) UBE3A/E6-AP mutations cause Angelman syndrome. Nat. Genet., 15, 70-73.
    • (1997) Nat. Genet. , vol.15 , pp. 70-73
    • Kishino, T.1    Lalande, M.2    Wagstaff, J.3
  • 10
    • 0028907874 scopus 로고
    • A family of proteins structurally and functionally related to the E6-AP ubiquitin-protein ligase
    • Huibregtse, J.M., Scheffner, M., Beaudenon, S. and Howley, P.M. (1995) A family of proteins structurally and functionally related to the E6-AP ubiquitin-protein ligase. Proc. Natl Acad. Sci. USA, 92, 2563-2567.
    • (1995) Proc. Natl Acad. Sci. USA , vol.92 , pp. 2563-2567
    • Huibregtse, J.M.1    Scheffner, M.2    Beaudenon, S.3    Howley, P.M.4
  • 11
    • 0027358723 scopus 로고
    • The HPV-16 E6 and E6-AP complex functions as a ubiquitin-protein ligase in the ubiquitination of p53
    • Scheffner, M., Huibregtse, J.M., Vierstra, R.D. and Howley, P.M. (1993) The HPV-16 E6 and E6-AP complex functions as a ubiquitin-protein ligase in the ubiquitination of p53. Cell, 75, 495-505.
    • (1993) Cell , vol.75 , pp. 495-505
    • Scheffner, M.1    Huibregtse, J.M.2    Vierstra, R.D.3    Howley, P.M.4
  • 12
    • 0028898424 scopus 로고
    • Protein ubiquitination involving an E1-E2-E3 enzyme ubiquitin thioester cascade
    • Scheffner, M., Nuber, U. and Huibregtse, J.M. (1995) Protein ubiquitination involving an E1-E2-E3 enzyme ubiquitin thioester cascade. Nature, 373, 81-83.
    • (1995) Nature , vol.373 , pp. 81-83
    • Scheffner, M.1    Nuber, U.2    Huibregtse, J.M.3
  • 13
    • 0032907106 scopus 로고    scopus 로고
    • The Angelman syndrome-associated protein, E6-AP, is a coactivator for the nuclear hormone receptor superfamily
    • Nawaz, Z., Lonard, D.M., Smith, C.L., Lev-Lehman, E., Tsai, S.Y., Tsai, M.J. and O'Malley, B.W. (1999) The Angelman syndrome-associated protein, E6-AP, is a coactivator for the nuclear hormone receptor superfamily. Mol. Cell Biol., 19, 1182-1189.
    • (1999) Mol. Cell Biol. , vol.19 , pp. 1182-1189
    • Nawaz, Z.1    Lonard, D.M.2    Smith, C.L.3    Lev-Lehman, E.4    Tsai, S.Y.5    Tsai, M.J.6    O'Malley, B.W.7
  • 14
    • 0032192481 scopus 로고    scopus 로고
    • Mutation of the Angelman ubiquitin ligase in mice causes increased cytoplasmic p53 and deficits of contextual learning and long-term potentiation
    • Jiang, Y.H., Armstrong, D., Albrecht, U., Atkins, C.M., Noebels, J.L., Eichele, G., Sweatt, J.D. andBeaudet, A.L. (1998)Mutation of the Angelman ubiquitin ligase in mice causes increased cytoplasmic p53 and deficits of contextual learning and long-term potentiation. Neuron, 21, 799-811.
    • (1998) Neuron , vol.21 , pp. 799-811
    • Jiang, Y.H.1    Armstrong, D.2    Albrecht, U.3    Atkins, C.M.4    Noebels, J.L.5    Eichele, G.6    Sweatt, J.D.7    Beaudet, A.L.8
  • 15
    • 0036197031 scopus 로고    scopus 로고
    • Neurobehavioral and electroencephalographic abnormalities inUbe3amaternal-deficientmice
    • Miura, K., Kishino, T., Li, E., Webber, H., Dikkes, P., Holmes, G.L. and Wagstaff, J. (2002) Neurobehavioral and electroencephalographic abnormalities inUbe3amaternal-deficientmice.Neurobiol.Dis., 9,149-159.
    • (2002) Neurobiol. Dis. , vol.9 , pp. 149-159
    • Miura, K.1    Kishino, T.2    Li, E.3    Webber, H.4    Dikkes, P.5    Holmes, G.L.6    Wagstaff, J.7
  • 16
    • 37549057995 scopus 로고    scopus 로고
    • The Angelman syndrome ubiquitin ligase localizes to the synapse and nucleus, and maternal deficiency results in abnormal dendritic spine morphology
    • Dindot, S.V., Antalffy, B.A., Bhattacharjee, M.B. and Beaudet, A.L. (2008) The Angelman syndrome ubiquitin ligase localizes to the synapse and nucleus, and maternal deficiency results in abnormal dendritic spine morphology. Hum. Mol. Genet., 17, 111-118.
    • (2008) Hum. Mol. Genet. , vol.17 , pp. 111-118
    • Dindot, S.V.1    Antalffy, B.A.2    Bhattacharjee, M.B.3    Beaudet, A.L.4
  • 18
    • 77950386683 scopus 로고    scopus 로고
    • Genomic imprinting of experience-dependent cortical plasticity by the ubiquitin ligase gene Ube3a
    • Sato, M. and Stryker, M.P. (2010) Genomic imprinting of experience-dependent cortical plasticity by the ubiquitin ligase gene Ube3a. Proc. Natl Acad. Sci. USA, 107, 5611-5616.
    • (2010) Proc. Natl Acad. Sci. USA , vol.107 , pp. 5611-5616
    • Sato, M.1    Stryker, M.P.2
  • 21
    • 4644295494 scopus 로고    scopus 로고
    • Biochemical analysis of Angelman syndrome-associated mutations in the E3 ubiquitin ligase E6-associated protein
    • Cooper, E.M., Hudson, A.W., Amos, J., Wagstaff, J. and Howley, P.M. (2004) Biochemical analysis of Angelman syndrome-associated mutations in the E3 ubiquitin ligase E6-associated protein. J. Biol. Chem., 279, 41208-41217.
    • (2004) J. Biol. Chem. , vol.279 , pp. 41208-41217
    • Cooper, E.M.1    Hudson, A.W.2    Amos, J.3    Wagstaff, J.4    Howley, P.M.5
  • 22
    • 0036135671 scopus 로고    scopus 로고
    • Genetic ablation of the steroid receptor coactivator-ubiquitin ligase, E6-AP, results in tissue-selective steroid hormone resistance and defects in reproduction
    • Smith, C.L., DeVera, D.G., Lamb, D.J., Nawaz, Z., Jiang, Y.H., Beaudet, A.L. and O'Malley, B.W. (2002) Genetic ablation of the steroid receptor coactivator-ubiquitin ligase, E6-AP, results in tissue-selective steroid hormone resistance and defects in reproduction.Mol. Cell Biol., 22, 525-535.
    • (2002) Mol. Cell Biol. , vol.22 , pp. 525-535
    • Smith, C.L.1    DeVera, D.G.2    Lamb, D.J.3    Nawaz, Z.4    Jiang, Y.H.5    Beaudet, A.L.6    O'Malley, B.W.7
  • 23
    • 33344469819 scopus 로고    scopus 로고
    • Multifunction steroid receptor coactivator, E6-associated protein, is involved in development of the prostate gland
    • Khan, O.Y., Fu, G., Ismail, A., Srinivasan, S., Cao, X., Tu, Y., Lu, S. and Nawaz, Z. (2006) Multifunction steroid receptor coactivator, E6-associated protein, is involved in development of the prostate gland. Mol. Endocrinol., 20, 544-559.
    • (2006) Mol. Endocrinol. , vol.20 , pp. 544-559
    • Khan, O.Y.1    Fu, G.2    Ismail, A.3    Srinivasan, S.4    Cao, X.5    Tu, Y.6    Lu, S.7    Nawaz, Z.8
  • 24
    • 60849109938 scopus 로고    scopus 로고
    • Chronic stress-induced hippocampal vulnerability: the glucocorticoid vulnerability hypothesis
    • Conrad, C.D. (2008) Chronic stress-induced hippocampal vulnerability: the glucocorticoid vulnerability hypothesis. Rev. Neurosci., 19, 395-411.
    • (2008) Rev. Neurosci. , vol.19 , pp. 395-411
    • Conrad, C.D.1
  • 25
    • 0031015517 scopus 로고    scopus 로고
    • Neurocircuitry of stress: central control of the hypothalamo-pituitary-adrenocortical axis
    • Herman, J.P. and Cullinan, W.E. (1997) Neurocircuitry of stress: central control of the hypothalamo-pituitary-adrenocortical axis. Trends Neurosci., 20, 78-84.
    • (1997) Trends Neurosci. , vol.20 , pp. 78-84
    • Herman, J.P.1    Cullinan, W.E.2
  • 26
  • 28
    • 34547689480 scopus 로고    scopus 로고
    • Chronic glucocorticoids increase hippocampal vulnerability to neurotoxicity under conditions that produce CA3 dendritic retraction but fail to impair spatial recognition memory
    • Conrad, C.D., McLaughlin, K.J., Harman, J.S., Foltz, C., Wieczorek, L., Lightner, E. and Wright, R.L. (2007) Chronic glucocorticoids increase hippocampal vulnerability to neurotoxicity under conditions that produce CA3 dendritic retraction but fail to impair spatial recognition memory. J. Neurosci., 27, 8278-8285.
    • (2007) J. Neurosci. , vol.27 , pp. 8278-8285
    • Conrad, C.D.1    McLaughlin, K.J.2    Harman, J.S.3    Foltz, C.4    Wieczorek, L.5    Lightner, E.6    Wright, R.L.7
  • 29
    • 0023691166 scopus 로고
    • A new one-trial test for neurobiological studies of memory in rats. 1: Behavioral data. Behav
    • Ennaceur, A. and Delacour, J. (1988) A new one-trial test for neurobiological studies of memory in rats. 1: Behavioral data. Behav. Brain Res., 31, 47-59.
    • (1988) Brain Res. , vol.31 , pp. 47-59
    • Ennaceur, A.1    Delacour, J.2
  • 30
    • 0027997328 scopus 로고
    • Comparison of different behavioral test situations used in psychopharmacology for measurement of anxiety
    • Belzung, C. and Le Pape, G. (1994) Comparison of different behavioral test situations used in psychopharmacology for measurement of anxiety. Physiol. Behav., 56, 623-628.
    • (1994) Physiol. Behav. , vol.56 , pp. 623-628
    • Belzung, C.1    Le Pape, G.2
  • 31
    • 77954851827 scopus 로고    scopus 로고
    • Defective GABAergic neurotransmission and pharmacological rescue of neuronal hyperexcitability in the amygdala in a mouse model of fragile X syndrome
    • Olmos-Serrano, J.L., Paluszkiewicz, S.M., Martin, B.S., Kaufmann, W.E., Corbin, J.G. and Huntsman, M.M. (2010) Defective GABAergic neurotransmission and pharmacological rescue of neuronal hyperexcitability in the amygdala in a mouse model of fragile X syndrome. J. Neurosci., 30, 9929-9938.
    • (2010) J. Neurosci. , vol.30 , pp. 9929-9938
    • Olmos-Serrano, J.L.1    Paluszkiewicz, S.M.2    Martin, B.S.3    Kaufmann, W.E.4    Corbin, J.G.5    Huntsman, M.M.6
  • 32
    • 24944475104 scopus 로고    scopus 로고
    • Anxiety, memory impairment, and locomotor dysfunction caused by a mutant thyroid hormone receptor alpha1 can be ameliorated by T3 treatment
    • Venero, C., Guadano-Ferraz, A., Herrero, A.I., Nordstrom, K., Manzano, J., de Escobar, G.M., Bernal, J. and Vennstrom, B. (2005) Anxiety, memory impairment, and locomotor dysfunction caused by a mutant thyroid hormone receptor alpha1 can be ameliorated by T3 treatment. Genes Dev., 19, 2152-2163.
    • (2005) Genes Dev. , vol.19 , pp. 2152-2163
    • Venero, C.1    Guadano-Ferraz, A.2    Herrero, A.I.3    Nordstrom, K.4    Manzano, J.5    de Escobar, G.M.6    Bernal, J.7    Vennstrom, B.8
  • 33
    • 0037439943 scopus 로고    scopus 로고
    • Genetic disruption of cortical interneuron development causes region- and GABA cell type-specific deficits, epilepsy, and behavioral dysfunction
    • Powell, E.M., Campbell, D.B., Stanwood, G.D., Davis, C., Noebels, J.L. and Levitt, P. (2003) Genetic disruption of cortical interneuron development causes region- and GABA cell type-specific deficits, epilepsy, and behavioral dysfunction. J. Neurosci., 23, 622-631.
    • (2003) J. Neurosci. , vol.23 , pp. 622-631
    • Powell, E.M.1    Campbell, D.B.2    Stanwood, G.D.3    Davis, C.4    Noebels, J.L.5    Levitt, P.6
  • 34
    • 11144267133 scopus 로고    scopus 로고
    • Chronic stress decreases the number of parvalbumin-immunoreactive interneurons in the hippocampus: prevention by treatment with a substance P receptor (NK1) antagonist
    • Czeh, B., Simon, M., van der Hart, M.G., Schmelting, B., Hesselink, M.B. and Fuchs, E. (2005) Chronic stress decreases the number of parvalbumin-immunoreactive interneurons in the hippocampus: prevention by treatment with a substance P receptor (NK1) antagonist. Neuropsychopharmacology, 30, 67-79.
    • (2005) Neuropsychopharmacology , vol.30 , pp. 67-79
    • Czeh, B.1    Simon, M.2    van der Hart, M.G.3    Schmelting, B.4    Hesselink, M.B.5    Fuchs, E.6
  • 35
    • 0041423730 scopus 로고    scopus 로고
    • Interneuron diversity series: rhythm and mood in perisomatic inhibition
    • Freund, T.F. (2003) Interneuron diversity series: rhythm and mood in perisomatic inhibition. Trends Neurosci., 26, 489-495.
    • (2003) Trends Neurosci. , vol.26 , pp. 489-495
    • Freund, T.F.1
  • 36
    • 33747816546 scopus 로고    scopus 로고
    • Lives and times of nuclear receptors
    • Alarid, E.T. (2006) Lives and times of nuclear receptors. Mol. Endocrinol., 20, 1972-1981.
    • (2006) Mol. Endocrinol. , vol.20 , pp. 1972-1981
    • Alarid, E.T.1
  • 37
    • 0000279559 scopus 로고    scopus 로고
    • Importance of the regulation of nuclear receptor degradation
    • Dennis, A.P., Haq, R.U. and Nawaz, Z. (2001) Importance of the regulation of nuclear receptor degradation. Front. Biosci., 6, D954-D959.
    • (2001) Front. Biosci. , vol.6
    • Dennis, A.P.1    Haq, R.U.2    Nawaz, Z.3
  • 38
    • 0036460384 scopus 로고    scopus 로고
    • Role of the paraventricular nucleus microenvironment in stress integration
    • Herman, J.P., Cullinan, W.E., Ziegler, D.R. and Tasker, J.G. (2002) Role of the paraventricular nucleus microenvironment in stress integration. Eur. J. Neurosci., 16, 381-385.
    • (2002) Eur. J. Neurosci. , vol.16 , pp. 381-385
    • Herman, J.P.1    Cullinan, W.E.2    Ziegler, D.R.3    Tasker, J.G.4
  • 40
    • 77953557625 scopus 로고    scopus 로고
    • Stress impairs GABAergic network function in the hippocampus by activating nongenomic glucocorticoid receptors and affecting the integrity of the parvalbumin-expressing neuronal network
    • Hu, W., Zhang, M., Czeh, B., Flugge, G. and Zhang, W. (2010) Stress impairs GABAergic network function in the hippocampus by activating nongenomic glucocorticoid receptors and affecting the integrity of the parvalbumin-expressing neuronal network. Neuropsychopharmacology, 35, 1693-1707.
    • (2010) Neuropsychopharmacology , vol.35 , pp. 1693-1707
    • Hu, W.1    Zhang, M.2    Czeh, B.3    Flugge, G.4    Zhang, W.5
  • 41
    • 0032912252 scopus 로고    scopus 로고
    • Stress and hippocampal plasticity
    • McEwen, B.S. (1999) Stress and hippocampal plasticity. Ann. Rev. Neurosci., 22, 105-122.
    • (1999) Ann. Rev. Neurosci. , vol.22 , pp. 105-122
    • McEwen, B.S.1
  • 42
    • 34447629382 scopus 로고    scopus 로고
    • Physiology and neurobiology of stress and adaptation: central role of the brain
    • McEwen, B.S. (2007) Physiology and neurobiology of stress and adaptation: central role of the brain. Physiol. Rev., 87, 873-904.
    • (2007) Physiol. Rev. , vol.87 , pp. 873-904
    • McEwen, B.S.1
  • 43
    • 0030935295 scopus 로고    scopus 로고
    • Cognitive dysfunction resulting from hippocampal hyperactivity-a possible cause of anxiety disorder? Pharmacol
    • McNaughton, N. (1997) Cognitive dysfunction resulting from hippocampal hyperactivity-a possible cause of anxiety disorder? Pharmacol. Biochem. Behav., 56, 603-611.
    • (1997) Biochem. Behav. , vol.56 , pp. 603-611
    • McNaughton, N.1
  • 44
    • 0036703905 scopus 로고    scopus 로고
    • Chronic stress induces contrasting patterns of dendritic remodeling in hippocampal and amygdaloid neurons
    • Vyas, A., Mitra, R., Shankaranarayana Rao, B.S. and Chattarji, S. (2002) Chronic stress induces contrasting patterns of dendritic remodeling in hippocampal and amygdaloid neurons. J. Neurosci., 22, 6810-6818.
    • (2002) J. Neurosci. , vol.22 , pp. 6810-6818
    • Vyas, A.1    Mitra, R.2    Shankaranarayana Rao, B.S.3    Chattarji, S.4
  • 45
    • 62349119948 scopus 로고    scopus 로고
    • Anxiety in children and adolescents with autism spectrum disorders
    • White, S.W., Oswald, D., Ollendick, T. and Scahill, L. (2009) Anxiety in children and adolescents with autism spectrum disorders. Clin. Psychol. Rev., 29, 216-229.
    • (2009) Clin. Psychol. Rev. , vol.29 , pp. 216-229
    • White, S.W.1    Oswald, D.2    Ollendick, T.3    Scahill, L.4
  • 46
    • 78651346564 scopus 로고    scopus 로고
    • Normal social seeking behavior, hypoactivity and reduced exploratory range in a mouse model of Angelman syndrome
    • Allensworth, M., Saha, A., Reiter, L.T. and Heck, D.H. (2011) Normal social seeking behavior, hypoactivity and reduced exploratory range in a mouse model of Angelman syndrome. BMC Genet., 12, 7.
    • (2011) BMC Genet. , vol.12 , pp. 7
    • Allensworth, M.1    Saha, A.2    Reiter, L.T.3    Heck, D.H.4
  • 47
    • 77957914925 scopus 로고    scopus 로고
    • Altered ultrasonic vocalization and impaired learning and memory in Angelman syndrome mouse model with a large maternal deletion from Ube3a to Gabrb3
    • Jiang, Y.H., Pan, Y., Zhu, L., Landa, L., Yoo, J., Spencer, C., Lorenzo, I., Brilliant, M., Noebels, J. and Beaudet, A.L. (2010) Altered ultrasonic vocalization and impaired learning and memory in Angelman syndrome mouse model with a large maternal deletion from Ube3a to Gabrb3. PLoS ONE, 5, e12278.
    • (2010) PLoS ONE , vol.5
    • Jiang, Y.H.1    Pan, Y.2    Zhu, L.3    Landa, L.4    Yoo, J.5    Spencer, C.6    Lorenzo, I.7    Brilliant, M.8    Noebels, J.9    Beaudet, A.L.10
  • 48
    • 47849125828 scopus 로고    scopus 로고
    • Behavior and neuropsychiatric manifestations in Angelman syndrome
    • Pelc, K., Cheron, G. and Dan, B. (2008) Behavior and neuropsychiatric manifestations in Angelman syndrome. Neuropsychiat. Dis. Treat., 4, 577-584.
    • (2008) Neuropsychiat. Dis. Treat. , vol.4 , pp. 577-584
    • Pelc, K.1    Cheron, G.2    Dan, B.3
  • 49
    • 0034945669 scopus 로고    scopus 로고
    • Angelman syndrome: evolution of the phenotype in adolescents and adults
    • Smith, J.C. (2001) Angelman syndrome: evolution of the phenotype in adolescents and adults. Develop. Med. Child Neurol., 43, 476-480.
    • (2001) Develop. Med. Child Neurol. , vol.43 , pp. 476-480
    • Smith, J.C.1
  • 51
    • 39749139563 scopus 로고    scopus 로고
    • Regulation of turnover of tumor suppressor p53 and cell growth by E6-AP, a ubiquitin protein ligase mutated in Angelman mental retardation syndrome
    • Mishra, A. and Jana, N.R. (2008) Regulation of turnover of tumor suppressor p53 and cell growth by E6-AP, a ubiquitin protein ligase mutated in Angelman mental retardation syndrome. Cell Mol. Life Sci., 65, 656-666.
    • (2008) Cell Mol. Life Sci. , vol.65 , pp. 656-666
    • Mishra, A.1    Jana, N.R.2
  • 52
    • 1642564603 scopus 로고    scopus 로고
    • Inhibition of proteasomal function by curcumin induces apoptosis through mitochondrial pathway
    • Jana, N.R., Dikshit, P., Goswami, A. and Nukina, N. (2004) Inhibition of proteasomal function by curcumin induces apoptosis through mitochondrial pathway. J. Biol. Chem., 279, 11680-11685.
    • (2004) J. Biol. Chem. , vol.279 , pp. 11680-11685
    • Jana, N.R.1    Dikshit, P.2    Goswami, A.3    Nukina, N.4
  • 53
    • 43149120469 scopus 로고    scopus 로고
    • E6-AP promotes misfolded polyglutamine proteins for proteasomal degradation and suppresses polyglutamine protein aggregation and toxicity
    • Mishra, A., Dikshit, P., Purkayastha, S., Sharma, J., Nukina, N. and Jana, N.R. (2008) E6-AP promotes misfolded polyglutamine proteins for proteasomal degradation and suppresses polyglutamine protein aggregation and toxicity. J. Biol. Chem., 283, 7648-7656.
    • (2008) J. Biol. Chem. , vol.283 , pp. 7648-7656
    • Mishra, A.1    Dikshit, P.2    Purkayastha, S.3    Sharma, J.4    Nukina, N.5    Jana, N.R.6
  • 54
    • 77956324028 scopus 로고    scopus 로고
    • Loss of dopaminergic neurons and resulting behavioural deficits in mouse model of Angelman syndrome
    • Mulherkar, S.A. and Jana, N.R. (2010) Loss of dopaminergic neurons and resulting behavioural deficits in mouse model of Angelman syndrome. Neurobiol. Dis., 40, 586-592.
    • (2010) Neurobiol. Dis. , vol.40 , pp. 586-592
    • Mulherkar, S.A.1    Jana, N.R.2


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