Polyomavirus Small T Antigen Induces Apoptosis in
Mammalian Cells through the UNC5B Pathway in a PP2A-
Dependent Manner
Sameer Ahmed Bhat,
a
Zarka Sarwar,
b
Syed Qaaifah Gillani,
b
Misbah Un Nisa,
b
Irfana Reshi,
a
Nusrat Nabi,
b
Shaozhen Xie,
c
Khalid M. Fazili,
a
Thomas M. Roberts,
c
Shaida Andrabi
b
a
Department of Biotechnology, University of Kashmir, Srinagar, India
b
Department of Biochemistry, University of Kashmir, Srinagar, India
c
Department of Cancer Biology, Dana Farber Cancer Institute, Harvard Medical School, Boston, Massachusetts, USA
ABSTRACT UNC5B is a dependence receptor that promotes survival in the pres-
ence of its ligand, netrin-1, while inducing cell death in its absence. The receptor
has an important role in the development of the nervous and vascular systems. It is
also involved in the normal turnover of intestinal epithelium. Netrin-1 and UNC5B
are deregulated in multiple cancers, including colorectal, neuroblastoma, and breast
tumors. However, the detailed mechanism of UNC5B function is not fully under-
stood. We have utilized the murine polyomavirus small T antigen (PyST) as a tool
to study UNC5B-mediated apoptosis. PyST is known to induce mitotic arrest fol-
lowed by extensive cell death in mammalian cells. Our results show that the ex-
pression of PyST increases mRNA levels of UNC5B by approximately 3-fold in os-
teosarcoma cells (U2OS) and also stabilizes UNC5B at the posttranslational level.
Furthermore, UNC5B is upregulated predominantly in those cells that undergo
mitotic arrest upon PyST expression. Interestingly, although its expression was
previously reported to be regulated by p53, our data show that the increase in
UNC5B levels by PyST is p53 independent. The posttranslational stabilization of
UNC5B by PyST is regulated by the interaction of PyST with PP2A. We also show
that netrin-1 expression, which is known to inhibit UNC5B apoptotic activity,
promotes survival of PyST-expressing cells. Our results thus suggest an important
role of UNC5B in small-T antigen-induced mitotic catastrophe that also requires
PP2A.
IMPORTANCE UNC5B, PP2A, and netrin-1 are deregulated in a variety of cancers.
UNC5B and PP2A are regarded as tumor suppressors, as they promote apoptosis
and are deleted or mutated in many cancers. In contrast, netrin-1 promotes survival
by inhibiting dependence receptors, including UNC5B, and is upregulated in many
cancers. Here, we show that UNC5B-mediated apoptosis can occur independently of
p53 but in a PP2A-dependent manner. A substantial percentage of cancers arise due
to p53 mutations and are insensitive to chemotherapeutic treatments that activate
p53. Unexpectedly, treatment of cancers having functional p53 with many conven-
tional drugs leads to the upregulation of netrin-1 through activated p53, which is
counterintuitive. Therefore, understanding the p53-independent mechanisms of the
netrin-UNC5B axis, such as those involving PP2A, assumes greater clinical signifi-
cance. Anticancer strategies utilizing anti-netrin-1 antibody treatment are already in
clinical trials.
KEYWORDS netrin 1, PP2A, polyomavirus, UNC5B, apoptosis, mitotic arrest, small T
antigen
Citation Bhat SA, Sarwar Z, Gillani SQ, Un Nisa
M, Reshi I, Nabi N, Xie S, Fazili KM, Roberts TM,
Andrabi S. 2020. Polyomavirus small T antigen
induces apoptosis in mammalian cells through
the UNC5B pathway in a PP2A-dependent
manner. J Virol 94:e02187-19. https://doi.org/
10.1128/JVI.02187-19.
Editor Lawrence Banks, International Centre
for Genetic Engineering and Biotechnology
Copyright © 2020 American Society for
Microbiology. All Rights Reserved.
Address correspondence to Shaida Andrabi,
shaida.andrabi@uok.edu.in.
Received 10 January 2020
Accepted 2 May 2020
Accepted manuscript posted online 13 May
2020
Published
TRANSFORMATION AND ONCOGENESIS
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