Aging Cell (2006) 5, pp331–344 Doi: 10.1111/j.1474-9726.2006.00223.x
© 2006 The Authors 331
Journal compilation © Blackwell Publishing Ltd/Anatomical Society of Great Britain and Ireland 2006
Blackwell Publishing Ltd
Age-specific hormonal decline is accompanied by
transcriptional changes in human sebocytes in vitro
Evgenia Makrantonaki,
1,3
James Adjaye,
2
Ralf Herwig,
2
Thore C. Brink,
2
Detlef Groth,
2
Claus Hultschig,
2
Hans Lehrach
2
and Christos C. Zouboulis
1,3
1
Laboratory for Biogerontology, Dermato-Pharmacology and
Dermato-Endocrinology, Institute of Clinical Pharmacology and
Toxicology, Charité Universitaetsmedizin Berlin, Campus Benjamin
Franklin, Hindenburgdamm 30, 12200 Berlin, Germany
2
Department of Vertebrate Genomics, Max Planck Institute for
Molecular Genetics, Ihnestrasse 73, 14195 Berlin, Germany
3
Departments of Dermatology and Immunology, Dessau Medical
Center, Auenweg 38, 06847 Dessau, Germany
Summary
The importance of hormones in endogenous aging has been
displayed by recent studies performed on animal models
and humans. To decipher the molecular mechanisms
involved in aging we maintained human sebocytes at
defined hormone-substituted conditions that corresponded
to average serum levels of females from 20 (f20) to 60
(f60) years of age. The corresponding hormone receptor
expression was demonstrated by reverse transcription–
polymerase chain reaction (RT–PCR), Western blotting and
immunocytochemistry. Cells at f60 produced significantly
lower lipids than at f20. Increased mRNA and protein
levels of c-Myc and increased protein levels of FN1, which
have been associated with aging, were detected in SZ95
sebocytes at f60 compared to those detected at f20 after
5 days of treatment. Expression profiling employing a cDNA
microarray composed of 15 529 cDNAs identified 899 genes
with altered expression levels at f20 vs. f60. Confirmation
of gene regulation was performed by real-time RT–PCR.
The functional annotation of these genes according to the
Gene Ontology identified pathways related to mitochondrial
function, oxidative stress, ubiquitin-mediated proteolysis,
cell cycle, immune responses, steroid biosynthesis and
phospholipid degradation – all hallmarks of aging. Twenty-
five genes in common with those identified in aging kidneys
and several genes involved in neurodegenerative diseases
were also detected. This is the first report describing the
transcriptome of human sebocytes and its modification by
a cocktail of hormones administered in age-specific levels
and provides an in vitro model system, which approximates
some of the hormone-dependent changes in gene transcrip-
tion that occur during aging in humans.
Key words: cDNA microarray; c-Myc; FN1; growth factors;
hormonal aging; neurodegenerative diseases, sebaceous
cells; sebaceous lipids; signaling pathways; steroids.
Introduction
Some of the theories regarding the generation of endogenous
aging support cellular senescence and decreased proliferative
ability (Hayflick, 1965; Smith & Pereira-Smith, 1996), reduction
of cellular DNA repair capacity, loss of telomeres with advancing
age (Allsopp et al., 1992; Bodnar et al., 1998), point mutations of
extranuclear mitochondrial DNA (Michikawa et al., 1999) which
may be associated with increased oxidative stress (Miquel, 1998)
and increased frequency of chromosomal abnormalities (Benn,
1976; Ly et al., 2000).
Complimentary evidence of the importance of hormones and
their metabolism on the aging process has come from recent
studies performed on animal models (Simon et al., 2003; Tatar
et al., 2003). This is not surprising as age-specific processes
in humans such as puberty and menopause are also controlled
hormonally. Serum levels of hormones decline with age in female
and male individuals (Roshan et al., 1999). This is due to
decreased functional reserve of the endocrine organs as a result
of aging per se or secondarily due to concurrent disease.
Skin, the largest organ of the body, usually provides the
first obvious evidence of the aging process and can be used as
a model for better understanding of the alterations occurring
with advancing age. Furthermore, as skin is the target for a
plethora of hormones and also a peripheral endocrine organ
in its own right (Zouboulis, 2000), it can be affected to a large
extent by the decline of hormones (Brincat, 2000).
During endogenous skin aging – observed in skin areas, which
are not in direct contact with environmental factors such as
ultraviolet (UV) irradiation – skin gradually loses its structural
and functional characteristics (Braverman & Fonferko, 1982;
Moragas et al., 1993; Ghadially et al., 1995; Fisher et al., 2002;
Raine-Fenning et al ., 2003; Wulf et al ., 2004). In particular,
morphological changes of the sebaceous gland cells and altera-
tions of their activity play a distinct role in the aging phenotype
of the skin. The size of sebaceous gland cells tends to decrease
with advancing age, while their number remains approximately
the same throughout life (Zouboulis & Boschnakow, 2001). Further-
more, sebaceous gland cells show an age-related reduced secretory
Evgenia Makrantonaki, James Adjaye and Ralf Herwig contributed equally to
this work.
Correspondence
Prof. Christos C. Zouboulis, Departments of Dermatology and
Immunology, Dessau Medical Center, Auenweg 38, 06847 Dessau,
Germany. Tel.: 49-340-5014000; fax: 49-340-5014025;
e-mail: christos.zouboulis@klinikum-dessau.de
Accepted for publication 13 May 2006