Schnabl et al
Hahn WC, Counter CM, Lundberg AS, Beijersbergen RL,
Brooks MW, and Weinberg RA (1999). Creation of human
tumour cells with defined genetic elements. Nature 400:464–
cancer-associated changes in human fibroblasts immortal-
ized with telomerase. Nat Genet 21:115–118.
Morin GB (1989). The human telomere terminal transferase
enzyme is a ribonucleoprotein that synthesizes TTAGGG
repeats. Cell 59:521–529.
468.
Harley CB, Futcher AB, and Greider CW (1990). Telomeres
shorten during ageing of human fibroblasts. Nature 345:458–
Moyzis RK, Buckingham JM, Cram LS, Dani M, Deaven LL,
Jones MD, Meyne J, Ratliff RL, and Wu JR (1988). A highly
conserved repetitive DNA sequence, (TTAGGG)n, present at
the telomeres of human chromosomes. Proc Natl Acad Sci
USA 85:6622–6626.
460.
Johnson LG, Mewshaw JP, Ni H, Friedmann T, Boucher RC,
and Olsen JC (1998). Effect of host modification and age on
airway epithelial gene transfer mediated by a murine leuke-
mia virus-derived vector. J Virol 72:8861–8872.
Murakami K, Abe T, Miyazawa M, Yamaguchi M, Masuda T,
Matsuura T, Nagamori S, Takeuchi K, Abe K, and Kyogoku M
(1995). Establishment of a new human cell line, LI90, exhib-
iting characteristics of hepatic Ito (fat-storing) cells. Lab
Invest 72:731–739.
Karlseder J, Broccoli D, Dai Y, Hardy S, and de Lange T
(1999). p53- and ATM-dependent apoptosis induced by
telomeres lacking TRF2. Science 283:1321–1325.
Kim NW, Piatyszek MA, Prowse KR, Harley CB, West MD, Ho
PL, Coviello GM, Wright WE, Weinrich SL, and Shay JW
Nakamura TM, Morin GB, Chapman KB, Weinrich SL, An-
drews WH, Lingner J, Harley CB, and Cech TR (1997).
Telomerase catalytic subunit homologs from fission yeast
and human. Science 277:955–959.
(1994). Specific association of human telomerase activity
with immortal cells and cancer. Science 266:2011–2015.
Kiyono T, Foster SA, Koop JI, McDougall JK, Galloway DA,
and Klingelhutz AJ (1998). Both Rb/p16INK4a inactivation
and telomerase activity are required to immortalize human
epithelial cells. Nature 396:84–88.
Neubauer K, Knittel T, Aurisch S, Fellmer P, and Ramadori G
(1996). Glial fibrillary acidic protein: A cell type specific
marker for Ito cells in vivo and in vitro. J Hepatol 24:719–730.
Knittel T, Kobold D, Saile B, Grundmann A, Neubauer K,
Piscaglia F, and Ramadori G (1999). Rat liver myofibroblasts
and hepatic stellate cells: Different cell populations of the
fibroblast lineage with fibrogenic potential. Gastroenterology
Niida H, Matsumoto T, Satoh H, Shiwa M, Tokutake Y,
Furuichi Y, and Shinkai Y (1998). Severe growth defect in
mouse cells lacking the telomerase RNA component. Nat
Genet 19:203–206.
1
17:1205–1221.
Nouchi T, Tanaka Y, Tsukada T, Sato C, and Marumo F
(1991). Appearance of alpha-smooth-muscle-actin-positive
cells in hepatic fibrosis. Liver 11:100–105.
Kojima H, Yokosuka O, Imazeki F, Saisho H, and Omata M
(
1997). Telomerase activity and telomere length in hepatocel-
lular carcinoma and chronic liver disease. Gastroenterology
12:493–500.
Ramadori G, Veit T, Schwogler S, Dienes HP, Knittel T,
Rieder H, and Meyer zum Buschenfelde KH (1990). Expres-
sion of the gene of the alpha-smooth muscle-actin isoform in
rat liver and in rat fat-storing (ITO) cells. Virchows Arch B Cell
Pathol Incl Mol Pathol 59:349–357.
1
Lang A, Schoonhoven R, Tuvia S, Brenner DA, and Rippe RA
2000). Nuclear factor kappaB in proliferation, activation, and
(
apoptosis in rat hepatic stellate cells. J Hepatol 33:49–58.
Lee HW, Blasco MA, Gottlieb GJ, Horner JW, 2nd, Greider
CW, and DePinho RA (1998). Essential role of mouse telom-
erase in highly proliferative organs. Nature 392:569–574.
Rippe RA, Almounajed G, and Brenner DA (1995). Sp1
binding activity increases in activated Ito cells. Hepatology
22:241–251.
Liu K, Schoonmaker MM, Levine BL, June CH, Hodes RJ,
and Weng NP (1999). Constitutive and regulated expression
of telomerase reverse transcriptase (hTERT) in human lym-
phocytes. Proc Natl Acad Sci USA 96:5147–5152.
Sahi J, Hamilton G, Sinz M, Barros S, Huang SM, Lesko LJ,
and LeCluyse EL (2000). Effect of troglitazone on cytochrome
P450 enzymes in primary cultures of human and rat hepato-
cytes. Xenobiotica 30:273–284.
Maher JJ, Bissell DM, Friedman SL, and Roll FJ (1988).
Collagen measured in primary cultures of normal rat hepato-
cytes derives from lipocytes within the monolayer. J Clin
Invest 82:450–459.
Schmitt-Graff A, Kruger S, Bochard F, Gabbiani G, and Denk
H (1991). Modulation of alpha smooth muscle actin and
desmin expression in perisinusoidal cells of normal and
diseased human livers. Am J Pathol 138:1233–1242.
Maher JJ and McGuire RF (1990). Extracellular matrix gene
expression increases preferentially in rat lipocytes and sinu-
soidal endothelial cells during hepatic fibrosis in vivo. J Clin
Invest 86:1641–1648.
Schnabl B, Kweon YO, Frederick JP, Wang XF, Rippe RA,
and Brenner DA (2001). The role of Smad3 in mediating
mouse hepatic stellate cell activation. Hepatology 34:89–
100.
Mak KM, Leo MA, and Lieber CS (1984). Alcoholic liver injury
in baboons: Transformation of lipocytes to transitional cells.
Gastroenterology 87:188–200.
Stefanovic B, Lindquist J, and Brenner DA (2000). The 5'
stem-loop regulates expression of collagen alpha1(I) mRNA
in mouse fibroblasts cultured in a three-dimensional matrix.
Nucleic Acids Res 28:641–647.
Meyerson M, Counter CM, Eaton EN, Ellisen LW, Steiner P,
Caddle SD, Ziaugra L, Beijersbergen RL, Davidoff MJ, Liu Q,
Bacchetti S, Haber DA, and Weinberg RA (1997). hEST2, the
putative human telomerase catalytic subunit gene, is up-
regulated in tumor cells and during immortalization. Cell
Takahara T, Kojima T, Miyabayashi C, Inoue K, Sasaki H,
Muragaki Y, and Ooshima A (1988). Collagen production in
fat-storing cells after carbon tetrachloride intoxication in the
rat: Immunoelectron microscopic observation of type I, type
III collagens, and prolyl hydroxylase. Lab Invest 59:509–521.
90:785–795.
Morales CP, Holt SE, Ouellette M, Kaur KJ, Yan Y, Wilson KS,
White MA, Wright WE, and Shay JW (1999). Absence of
Vaziri H and Benchimol S (1998). Reconstitution of telomer-
ase activity in normal human cells leads to elongation of
3
32 Laboratory Investigation • March 2002 • Volume 82 • Number 3