Cell and Tissue Research

, Volume 341, Issue 3, pp 341–347 | Cite as

Aging of cerebellar Purkinje cells



Cerebellar Purkinje cells (PCs), the sole output neurons in the cerebellar cortex, play an important role in the cerebellar circuit. PCs appear to be rather sensitive to aging, exhibiting significant changes in both morphology and function during senescence. This article reviews such changes during the normal aging process, including a decrease in the quantity of cells, atrophy in the soma, retraction in the dendritic arborizations, degeneration in the subcellular organelles, a decline in synapse density, disorder in the neurotransmitter system, and alterations in electrophysiological properties. Although these deteriorative changes occur during aging, compensatory mechanisms exist to counteract the impairments in the aging PCs. The possible neural mechanisms underlying these changes and potential preventive treatments are discussed.


Cerebellum Purkinje cells Aging Morphology Function 


  1. Alcock J, Scotting P, Sottile V (2007) Bergmann glia as putative stem cells of the mature cerebellum. Med Hypotheses 69:341–345CrossRefPubMedGoogle Scholar
  2. Andersen BB, Gundersen HJ, Pakkenberg B (2003) Aging of the human cerebellum: a stereological study. J Comp Neurol 466:356–365CrossRefPubMedGoogle Scholar
  3. Atamna H (2004) Heme, iron, and the mitochondrial decay of ageing. Ageing Res Rev 3:303–318CrossRefPubMedGoogle Scholar
  4. Bakalian A, Corman B, Delhaye-Bouchaud N, Mariani J (1991) Quantitative analysis of the Purkinje cell population during extreme ageing in the cerebellum of the Wistar/Louvain rat. Neurobiol Aging 12:425–430CrossRefPubMedGoogle Scholar
  5. Bertoni-Freddari C, Giuli C, Pieri C, Paci D (1986) Age-related morphological rearrangements of synaptic junctions in the rat cerebellum and hippocampus. Arch Gerontol Geriatr 5:297–304CrossRefPubMedGoogle Scholar
  6. Bertoni-Freddari C, Fattoretti P, Paoloni R, Caselli U, Galeazzi L, Meier-Ruge W (1996) Synaptic structural dynamics and aging. Gerontology 42:170–180CrossRefPubMedGoogle Scholar
  7. Bickford P (1993) Motor learning deficits in aged rats are correlated with loss of cerebellar noradrenergic function. Brain Res 620:133–138CrossRefPubMedGoogle Scholar
  8. Bickford PC, Hoffer BJ, Freedman R (1985) Interaction of norepinephrine with Purkinje cell responses to cerebellar afferent inputs in aged rats. Neurobiol Aging 6:89–94CrossRefPubMedGoogle Scholar
  9. Bickford PC, Shukitt-Hale B, Joseph J (1999) Effects of aging on cerebellar noradrenergic function and motor learning: nutritional interventions. Mech Ageing Dev 111:141–154CrossRefPubMedGoogle Scholar
  10. Bickford-Wimer PC, Granholm AC, Gerhardt GA (1988) Cerebellar noradrenergic systems in aging: studies in situ and in in oculo grafts. Neurobiol Aging 9:591–599CrossRefPubMedGoogle Scholar
  11. Binder DK (2007) Neurotrophins in the dentate gyrus. Prog Brain Res 163:371–397CrossRefPubMedGoogle Scholar
  12. Binstock RH, Fishman JR, Juengst ET (2006) Boundaries and labels: anti-aging medicine and science. Rejuvenation Res 9:433–435CrossRefPubMedGoogle Scholar
  13. Caston J, Hilber P, Chianale C, Mariani J (2003) Effect of training on motor abilities of heterozygous staggerer mutant (Rora(+)/Rora(sg)) mice during aging. Behav Brain Res 141:35–42CrossRefPubMedGoogle Scholar
  14. Caston J, Chianale C, Mariani J (2004) Spatial memory of heterozygous staggerer (Rora(+)/Rora(sg)) versus normal (Rora(+)/Rora(+)) mice during aging. Behav Genet 34:319–324CrossRefPubMedGoogle Scholar
  15. Chae CH, Kim HT (2009) Forced, moderate-intensity treadmill exercise suppresses apoptosis by increasing the level of NGF and stimulating phosphatidylinositol 3-kinase signaling in the hippocampus of induced aging rats. Neurochem Int 55:208–213CrossRefPubMedGoogle Scholar
  16. Chen S, Hillman DE (1999) Dying-back of Purkinje cell dendrites with synapse loss in aging rats. J Neurocytol 28:187–196CrossRefPubMedGoogle Scholar
  17. Chintala S, Novak EK, Spernyak JA, Mazurchuk R, Torres G, Patel S, Busch K, Meeder BA, Horowitz JM, Vaughan MM, Swank RT (2009) The Vps33a gene regulates behavior and cerebellar Purkinje cell number. Brain Res 1266:18–28CrossRefPubMedGoogle Scholar
  18. Chung YH, Shin C, Park KH, Cha CI (2000) Immunohistochemical study on the distribution of neuronal voltage-gated calcium channels in the rat cerebellum. Brain Res 865:278–282CrossRefPubMedGoogle Scholar
  19. Chung YH, Shin CM, Kim MJ, Lee BK, Cha CI (2001a) Age-related changes in the distribution of Kv1.1, Kv1.2 channel subunits in the rat cerebellum. Brain Res 897:193–198CrossRefPubMedGoogle Scholar
  20. Chung YH, Shin CM, Kim MJ, Shin DH, Yoo YB, Cha CI (2001b) Differential alterations in the distribution of voltage-gated calcium channels in aged rat cerebellum. Brain Res 903:247–252CrossRefPubMedGoogle Scholar
  21. Chung YH, Shin CM, Joo KM, Kim MJ, Cha CI (2002) Age-related upregulation of insulin-like growth factor receptor type I in rat cerebellum. Neurosci Lett 330:65–68CrossRefPubMedGoogle Scholar
  22. Chung YH, Joo KM, Kim MJ, Cha CI (2003) Age-related changes in the distribution of Na(v)1.1 and Na(v)1.2 in rat cerebellum. NeuroReport 14:841–845CrossRefPubMedGoogle Scholar
  23. Dhar P, Mohari N, Mehra RD (2007) Preliminary morphological and morphometric study of rat cerebellum following sodium arsenite exposure during rapid brain growth (RBG) period. Toxicology 234:10–20CrossRefPubMedGoogle Scholar
  24. Dlugos C (2005) Analyses of smooth endoplasmic reticulum of cerebellar parallel fibers in aging, ethanol-fed rats. Alcohol 35:67–73CrossRefPubMedGoogle Scholar
  25. Dlugos CA, Pentney RJ (1994) Morphometric analyses of Purkinje and granule cells in aging F344 rats. Neurobiol Aging 15:435–440CrossRefPubMedGoogle Scholar
  26. Drüge H, Heinsen H, Heinsen YL (1986) Quantitative studies in ageing Chbb: THOM (Wistar) rats. II. Neuron numbers in lobules I, VIb+c and X. Bibl Anat 28:121–137PubMedGoogle Scholar
  27. Erickson KI, Prakash RS, Voss MW, Chaddock L, Heo S, McLaren M, Pence BD, Martin SA, Vieira VJ, Woods JA, McAuley E, Kramer AF (2010) Brain-derived neurotrophic factor is associated with age-related decline in hippocampal volume. J Neurosci 30:5368–5375CrossRefPubMedGoogle Scholar
  28. Fattoretti P, Bertoni-Freddari C, Caselli U, Paoloni R, Meier-Ruge W (1996) Morphologic changes in cerebellar mitochondria during aging. Anal Quant Cytol Histol 18:205–208PubMedGoogle Scholar
  29. Fattoretti P, Bertoni-Freddari C, Caselli U, Paoloni R, Meier-Ruge W (1998) Impaired succinic dehydrogenase activity of rat Purkinje cell mitochondria during aging. Mech Ageing Dev 101:175–182CrossRefPubMedGoogle Scholar
  30. Felici L, Bronzetti E, Amenta F (1989) Enzyme histochemistry of glutamate dehydrogenase in ageing rat cerebellar cortex. Mech Ageing Dev 47:199–205CrossRefPubMedGoogle Scholar
  31. George O, Vallée M, Le Moal M, Mayo W (2006) Neurosteroids and cholinergic systems: implications for sleep and cognitive processes and potential role of age-related changes. Psychopharmacology 186:402–413CrossRefPubMedGoogle Scholar
  32. Glick R, Bondareff W (1979) Loss of synapses in the cerebellar cortex of the senescent rat. J Gerontol 34:818–822PubMedGoogle Scholar
  33. Gorbunova V, Seluanov A, Mao Z, Hine C (2007) Changes in DNA repair during aging. Nucleic Acids Res 35:7466–7474CrossRefPubMedGoogle Scholar
  34. Goukassian DA, Bagheri S, Keeb L, Eller MS, Gilchrest BA (2002) DNA oligonucleotide treatment corrects the age-associated decline in DNA repair capacity. FASEB J 16:754–756PubMedGoogle Scholar
  35. Gruart A, Muñoz MD, Delgado-García JM (2006) Involvement of the CA3-CA1 synapse in the acquisition of associative learning in behaving mice. J Neurosci 26:1077–1087CrossRefPubMedGoogle Scholar
  36. Hadj-Sahraoui N, Frederic F, Zanjani H, Delhaye-Bouchaud N, Herrup K, Mariani J (2001) Progressive atrophy of cerebellar PC dendrites during aging of the heterozygous staggerer mouse (Rora+/sg). Dev Brain Res 126:201–209CrossRefGoogle Scholar
  37. Hall TC, Millerakh Corsellis JAN (1975) Variations in the human Purkinje cell population according to age and sex. Neuropathol Appl Neurobiol 1:267–292CrossRefGoogle Scholar
  38. Hilber P, Caston J (2001) Motor skills and motor learning in Lurcher mutant mice during aging. Neuroscience 102:615–623CrossRefPubMedGoogle Scholar
  39. Ho YS, Yu MS, Yik SY, So KF, Yuen WH, Chang RC (2009) Polysaccharides from Wolfberry Antagonizes Glutamate Excitotoxicity in Rat Cortical Neurons. Cell Mol Neurobiol 29:1233–1244CrossRefPubMedGoogle Scholar
  40. Hogan MJ (2004) The cerebellum in thought and action: a fronto-cerebellar aging hypothesis. New Ideas in Psychol 22:97–125CrossRefGoogle Scholar
  41. Hua T, Kao C, Sun Q, Li X, Zhou Y (2008) Decreased proportion of GABA neurons accompanies age-related degradation of neuronal function in cat striate cortex. Brain Res Bull 75:119–125CrossRefPubMedGoogle Scholar
  42. Huang CM, Brown N, Huang RH (1999) Age-related changes in the cerebellum: parallel fibers. Brain Res 840:148–152CrossRefPubMedGoogle Scholar
  43. Huang CM, Miyamoto H, Huang RH (2006a) The mouse cerebellum from 1 to 34 months: parallel fibers. Neurobiol Aging 27:1715–1718CrossRefPubMedGoogle Scholar
  44. Huang CM, Wang L, Huang RH (2006b) Cerebellar granule cell: ascending axon and parallel fiber. Eur J Neurosci 23:1731–1737CrossRefPubMedGoogle Scholar
  45. Jankovski A, Rossi F, Sotelo C (1996) Neuronal precursors in the postnatal mouse cerebellum are fully committed cells: evidence from heterochronic transplantations. Eur J Neurosci 8:2308–2319CrossRefPubMedGoogle Scholar
  46. Jin K, Galvan V (2007) Endogenous neural stem cells in the adult brain. J Neuroimmune Pharmacol 2:236–242CrossRefPubMedGoogle Scholar
  47. Joseph JA, Shukitt-Hale B, Denisova NA, Prior RL, Cao G, Martin A, Taglialatela G, Bickford PC (1998) Long-term dietary strawberry, spinach, or vitamin E supplementation retards the onset of age-related neuronal signal-transduction and cognitive behavioral deficits. J Neurosci 18:8047–8055PubMedGoogle Scholar
  48. Klein C, Butt SJ, Machold RP, Johnson JE, Fishell G (2005) Cerebellum- and forebrain-derived stem cells possess intrinsic regional character. Development 132:4497–4508CrossRefPubMedGoogle Scholar
  49. Kodama T, Itsukaichi-Nishida Y, Fukazawa Y, Wakamori M, Miyata M, Molnar E, Mori Y, Shigemoto R, Imoto K (2006) A CaV2.1 calcium channel mutation rocker reduces the number of postsynaptic AMPA receptors in parallel fiber–Purkinje cell synapses. Eur J Neurosci 24:2993–3007CrossRefPubMedGoogle Scholar
  50. Lärkfors L, Lindsay RM, Alderson RF (1994) Ciliary neurotrophic factor enhances the survival of Purkinje cells in vitro. Eur J Neurosci 6:1015–1025CrossRefPubMedGoogle Scholar
  51. Larsen JO, Skalicky M, Viidik A (2000) Does long-term physical exercise counteract age-related Purkinje cell loss? A stereological study of rat cerebellum. Comp Neurol 428:213–222CrossRefGoogle Scholar
  52. Lee JY, Lyoo IK, Kim SU, Jang HS, Lee DW, Jeon HJ, Park SC, Cho MJ (2005) Intellect declines in healthy elderly subjects and cerebellum. Psychiatry Clin Neurosci 59:45–51CrossRefPubMedGoogle Scholar
  53. Lee KJ, Jung JG, Arii T, Imoto K, Rhyu IJ (2007) Morphological changes in dendritic spines of Purkinje cells associated with motor learning. Neurobiol Learn Mem 88:445–450CrossRefPubMedGoogle Scholar
  54. Louis ED, Faust PL, Vonsattel JP, Honig LS, Henchcliffe C, Pahwa R, Lyons KE, Rios E, Erickson-Davis C, Moskowitz CB, Lawton A (2009) Older onset essential tremor: more rapid progression and more degenerative pathology. Mov Disord 24:1606–1612CrossRefPubMedGoogle Scholar
  55. Manev H, Uz T, Sugaya K, Qu T (2000) Putative role of neuronal 5-lipoxygenase in an aging brain. FASEB J 14:1464–1469CrossRefPubMedGoogle Scholar
  56. Markowska AL, Mooney M, Sonntag WE (1998) Insulin-like growth factor-1 ameliorates age-related behavioral deficits. Neuroscience 87:559–569CrossRefPubMedGoogle Scholar
  57. Martinez Gagliardo K, Clebis NK, Stabille SR, De Britto MR, De Sousa JM, De Souza RR (2008) Exercise reduces inhibitory neuroactivity and protects myenteric neurons from age-related neurodegeneration. Auton Neurosci 141:31–37CrossRefPubMedGoogle Scholar
  58. Marwaha J, Hoffer B, Pittman R, Freedman R (1980) Age-related electrophysiological changes in rat cerebellum. Brain Res 201:85–97CrossRefPubMedGoogle Scholar
  59. Mattay VS, Fera F, Tessitore A, Hariri AR, Das S, Callicott JH, Weinberger DR (2002) Neurophysiological correlates of age-related changes in human motor function. Neurology 58:630–635PubMedGoogle Scholar
  60. Monteiro RA (1991) Age-related quantitative changes in the organelles of rat neocerebellar Purkinje cells. Histol Histopathol 6:9–20PubMedGoogle Scholar
  61. Monteiro RA, Rocha E, Marini-Abreu MM (1992) Age-related quantitative changes in inhibitory axo-somatic synapses on Purkinje cells of rat neocerebellum (Crus I and Crus II). J Submicrosc Cytol Pathol 24:351–357PubMedGoogle Scholar
  62. Monteiro RA, Rocha E, Marini-Abreu MM (1994) Heterogeneity and death of Purkinje cells of rat neocerebellum (Crus I and Crus II): hypothetic mechanisms based on qualitative and quantitative microscopical data. J Hirnforsch 35:205–222PubMedGoogle Scholar
  63. Monteiro RA, Henrique RM, Rocha E, Silva MW, Oliveira MH (2000) Quantitative age-changes in endoplasmic reticulum and nucleus of cerebellar granule cells. Neurobiol Aging 21:97–105CrossRefPubMedGoogle Scholar
  64. Niblock MM, Brunso-Bechtold JK, Riddle DR (2000) Insulin-like growth factor I stimulates dendritic growth in primary somatosensory cortex. J Neurosci 20:4165–4176PubMedGoogle Scholar
  65. Nosal G (1979) Neuronal involution during ageing. Ultrastructural study in the rat cerebellum. Mech Ageing Dev 10:295–314CrossRefPubMedGoogle Scholar
  66. Ogata R, Ikari K, Hayashi M, Tamai K, Tagawa K (1984) Age-related changes in the Purkinje's cells in the rat cerebellar cortex: a quantitative electron microscopic study. Folia Psychiatr Neurol Jpn 38:159–167PubMedGoogle Scholar
  67. Ohyama H, Hiramatsu M, Ogawa N, Mori A (1995) Age-related differences in synaptosomal membrane fluidity. Biochem Mol Biol Int 37:133–140PubMedGoogle Scholar
  68. Ojaimi J, Masters CL, Opeskin K, McKelvie P, Byrne E (1999) Mitochondrial respiratory chain activity in the human brain as a function of age. Mech Ageing Dev 111:39–47CrossRefPubMedGoogle Scholar
  69. Parfitt KD (1988) Age-related electrophysiological changes in cerebellar noradrenergic receptors. Age 11:120–127CrossRefGoogle Scholar
  70. Patrick GW, Anderson WJ (2000) Dendritic alterations of cerebellar Purkinje neurons in postnatally lead-exposed kittens. Dev Neurosci 22:320–328CrossRefPubMedGoogle Scholar
  71. Paul R, Grieve SM, Chaudary B, Gordon N, Lawrence J, Cooper N, Clark CR, Kukla M, Mulligan R, Gordon E (2009) Relative contributions of the cerebellar vermis and prefrontal lobe volumes on cognitive function across the adult lifespan. Neurobiol Aging 30:457–465CrossRefPubMedGoogle Scholar
  72. Pentney RJ, Mullan BA, Felong AM, Dlugos CA (2002) The total numbers of cerebellar granule neurons in young and aged Fischer 344 and Wistar–Kyoto rats do not change as a result of lengthy ethanol treatment. Cerebellum 1:79–89CrossRefPubMedGoogle Scholar
  73. Pires RS, Real CC, Folador TS, Tellini NR, Torrão AS, Britto LR (2010) Differential response of AMPA and NMDA glutamate receptors of Purkinje cells to aging of the chicken cerebellum. Neurosci Lett 478:146–149Google Scholar
  74. Poe BH, Linville C, Brunso-Bechtold J (2001) Age-related decline of presumptive inhibitory synapses in the sensorimotor cortex as revealed by the physical disector. J Comp Neurol 439:65–72CrossRefPubMedGoogle Scholar
  75. Porras-García E, Cendelin J, Domínguez-del-Toro E, Vozeh F, Delgado-García JM (2005) Purkinje cell loss affects differentially the execution, acquisition and prepulse inhibition of skeletal and facial motor responses in Lurcher mice. Eur J Neurosci 21:979–988CrossRefPubMedGoogle Scholar
  76. Porras-García E, Sánchez-Campusano R, Matínez-Vargas D, Domínguez-Del-Toro E, Cendelín J, Vozeh F, Delgado-Garcia JM (2010) Behavioral characteristics, associative learning capabilities, and dynamic association mapping in an animal model of cerebellar degeneration. J Neurophysiol 104:346–365Google Scholar
  77. Quackenbush LJ, Ngo H, Pentney RJ (1990) Evidence for nonrandom regression of dendrites of Purkinje neurons during aging. Neurobiol Aging 11:111–115CrossRefPubMedGoogle Scholar
  78. Rao SM, Mattson PM (2001) Stem cells and aging: expanding the possibilities. Mech Ageing Dev 122:713–734CrossRefPubMedGoogle Scholar
  79. Rogers J, Zornetzer SF, Bloom FE (1981) Senescent pathology of cerebellum: Purkinje neurons and their parallel fiber afferents. Neurobiol Aging 2:15–25CrossRefPubMedGoogle Scholar
  80. Rogers J, Zornetzer SF, Bloom FE, Mervis RE (1984) Senescent microstructural changes in rat cerebellum. Brain Res 292:23–32CrossRefPubMedGoogle Scholar
  81. Sabbatini M, Barili P, Bronzetti E, Zaccheo D, Amenta F (1999) Age-related changes of glial fibrillary acidic protein immunoreactive astrocytes in the rat cerebellar cortex. Mech Ageing Dev 108:165–172CrossRefPubMedGoogle Scholar
  82. Schaller KL, Caldwell JH (2003) Expression and distribution of voltage-gated sodium channels in the cerebellum. Cerebellum 2:2–9CrossRefPubMedGoogle Scholar
  83. Schumacher M, Weill-Engerer S, Liere P, Robert F, Franklin RJ, Garcia-Segura LM, Lambert JJ, Mayo W, Melcangi RC, Parducz A, Suter U, Carelli C, Baulieu EE, Akwa Y (2003) Steroid hormones and neurosteroids in normal and pathological aging of the nervous system. Prog Neurobiol 71:3–29CrossRefPubMedGoogle Scholar
  84. Seo MY, Chung SY, Choi WK, Seo YK, Jung SH, Park JM, Seo MJ, Park JK, Kim JW, Park CS (2009) Anti-aging effect of rice wine in cultured human fibroblasts and keratinocytes. J Biosci Bioeng 107:266–271CrossRefPubMedGoogle Scholar
  85. Servais L, Hourez R, Bearzatto B, Gall D, Schiffmann SN, Cheron G (2007) Purkinje cell dysfunction and alteration of long-term synaptic plasticity in fetal alcohol syndrome. Proc Natl Acad Sci USA 104:9858–9863CrossRefPubMedGoogle Scholar
  86. Sjöbeck M, Englund E (2001) Alzheimer's disease and the cerebellum: a morphologic study on neuronal and glial changes. Dement Geriatr Cogn Disord 12:211–218CrossRefPubMedGoogle Scholar
  87. Sun Y, Jin K, Mao XO, Xie L, Peel A, Childs JT, Logvinova A, Wang X, Greenberg DA (2005) Effect of aging on neuroglobin expression in rodent brain. Neurobiol Aging 26:275–278CrossRefPubMedGoogle Scholar
  88. Takahashi E, Niimi K, Itakura C (2009) Motor coordination impairment in aged heterozygous rolling Nagoya, Cav2.1 mutant mice. Brain Res 1279:50–57CrossRefPubMedGoogle Scholar
  89. Taniwaki T, Okayama A, Yoshiura T, Togao O, Nakamura Y, Yamasaki T, Ogata K, Shigeto H, Ohyagi Y, Kira J, Tobimatsu S (2007) Age-related alterations of the functional interactions within the basal ganglia and cerebellar motor loops in vivo. Neuroimage 36:1263–1276CrossRefPubMedGoogle Scholar
  90. Tolbert DL, Clark BR (2003) GDNF and IGF-I trophic factors delay hereditary Purkinje cell degeneration and the progression of gait ataxia. Exp Neurol 183:205–219CrossRefPubMedGoogle Scholar
  91. Tranquilli Leali FM, Artico M, Potenza S, Cavallotti C (2003) Age-related changes of monoaminooxidases in rat cerebellar cortex. Eur J Histochem 47:81–86PubMedGoogle Scholar
  92. Tu PH, Robinson KA, de Snoo F, Eyer J, Peterson A, Lee VM, Trojanowski JQ (1997) Selective degeneration of Purkinje cells with Lewy body-like inclusions in aged NFHLACZ transgenic mice. J Neurosci 17:1064–1074PubMedGoogle Scholar
  93. von Bohlen und Halbach O, Unsicker K (2002) Morphological alterations in the amygdala and hippocampus of mice during ageing. Eur J Neurosci 16:2434–2440CrossRefPubMedGoogle Scholar
  94. Woodruff-Pak DS (2006) Stereological estimation of Purkinje neuron number in C57BL/6 mice and its relation to associative learning. Neuroscience 141:233–243CrossRefPubMedGoogle Scholar
  95. Woodruff-Pak DS, Vogel RW 3rd, Ewers M, Coffey J, Boyko OB, Lemieux SK (2001) MRI-assessed volume of cerebellum correlates with associative learning. Neurobiol Learn Mem 76:342–357CrossRefPubMedGoogle Scholar
  96. Woodruff-Pak DS, Foy MR, Akopian GG, Lee KH, Zach J, Nguyen KP, Comalli DM, Kennard JA, Agelan A, Thompson RF (2010) Differential effects and rates of normal aging in cerebellum and hippocampus. Proc Natl Acad Sci USA 107:1624–1629CrossRefPubMedGoogle Scholar
  97. Zhang CZ, Hua TM, Zhu ZM, Luo X (2006) Age-related changes of structures in cerebellar cortex of cat. J Biosci 31:55–60CrossRefPubMedGoogle Scholar

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© Springer-Verlag 2010

Authors and Affiliations

  1. 1.Department of Biological Science and Technology/State Key Laboratory of Pharmaceutical BiotechnologySchool of Life Sciences, Nanjing UniversityNanjingChina
  2. 2.School of Life SciencesAnqing Teachers CollegeAnqingPeople’s Republic of China
  3. 3.School of Life SciencesAnhui Normal UniversityWuhuPeople’s Republic of China

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