Cellular Microbiology (2004) 6(8), 743–751 doi:10.1111/j.1462-5822.2004.00400.x
© 2004 Blackwell Publishing Ltd
Blackwell Science, LtdOxford, UKCMICellular Microbiology 1462-5814Blackwell Publishing Ltd, 20046 8743751Original ArticleJ. Park et al.AnkA binds to host DNA and nuclear proteins
Received 22 December, 2003; revised 27 February, 2004; accepted
2 March, 2004. *For correspondence. E-mail sdumler@jhmi.edu; Tel.
(+1) 410 955 8654; Fax (+1) 443 287 3665. †Present address:
Department of Veterinary Internal Medicine, College of Veterinary
Medicine, Chonbuk National University, Jeonju, Jeonbuk 561–756,
Korea.
Anaplasma phagocytophilum AnkA binds to granulocyte
DNA and nuclear proteins
Jinho Park,
1†
Kee Jun Kim,
2
Kyoung-seong Choi,
1
Dennis J. Grab
2
and J. Stephen Dumler
1
*
1
Division of Medical Microbiology, Department of
Pathology, The Johns Hopkins University School of
Medicine, Ross Research Building, 720 Rutland Avenue,
Baltimore, Maryland, USA.
2
Division of Infectious Diseases, Department of Pediatrics,
The Johns Hopkins University School of Medicine,
Baltimore, Maryland, USA.
Summary
Human granulocytic anaplasmosis (HGA) is caused
by the obligate intracellular bacterium Anaplasma
phagocytophilum. The bacterium infects, survives,
propagates in, and alters neutrophil phenotype, indi-
cating unique survival mechanisms. AnkA is the only
known A. phagocytophilum component that gains
access beyond neutrophil vacuoles and is trans-
ported to the infected host cell nucleus. The ability of
native and recombinant AnkA to bind DNA and
nuclear proteins from host HL-60 cells was assessed
by the use of immunoprecipitation after cis-
diamminedichloroplatinum (cis-DDP) DNA-protein
crosslinking, by probing uninfected HL-60 cell nuclear
lysates for AnkA binding, and by recovery and
sequence analysis of immunoprecipitated DNA. AnkA
binds HL-60 cell DNA as well as nuclear proteins of
approximately 86, 53 and 25 kDa, whereas recombi-
nant A. phagocytophilum Msp2 or control proteins do
not. DNA immunoprecipitation reveals AnkA binding
to a variety of target genes in the human genome,
including genes that encode proteins with ATPase,
tyrosine phosphatase and NADH dehydrogenase-like
functions. These data indicate that AnkA could exert
some effect on cells through binding to protein:DNA
complexes in neutrophil nuclei. Whether AnkA bind-
ing leads to neutrophil functional alterations, and how
such alterations might occur will depend upon defin-
itive identification of binding partners and associated
metabolic and biochemical pathways.
Introduction
Anaplasma phagocytophilum is a tick-borne, obligate
intracellular bacterium that infects neutrophils of mam-
mals, including man (Dumler et al., 2001). The unique
niche of this bacterium implies that it has developed spe-
cialized systems for survival and propagation. It is now
well documented that A. phagocytophilum alters neutro-
phil function by inducing an ‘activated-deactivated’ pheno-
type, wherein infected neutrophils are incapable of
generating phagocyte oxidase, are defective at adhering
to and transmigrating across activated endothelium, and
are defective at phagocytosis and microbicidal activity. Yet,
infection activates cells for production of proinflammatory
responses with chemokine secretion, degranulation and
release of metalloproteases (Woldehiwet, 1987; Banerjee
et al., 2000; Klein et al., 2000; Choi et al., 2003; Park
et al., 2003a). The mechanisms that underlie control of
these various cellular processes by the bacterium are
increasingly studied and appear to involve regulation at
the level of host cell mRNA, via signal transduction, and
perhaps by effects upon transcription factors (Banerjee
et al., 2000; Carlyon et al., 2002; Kim and Rikihisa, 2002;
Lin and Rikihisa, 2003).
AnkA (also known as Ank) is a unique protein of A.
phagocytophilum that is high molecular weight (160 kDa),
not associated with the bacterial membrane, but is found
localized within nuclei of infected granulocyte hosts, pre-
sumably after secretion and transport through at least
three distinct membranes (Caturegli et al., 2000).
Although its close ultrastructural association with con-
densed chromatin and the presence of multiple ankyrin
repeat domains tempts speculation about a role in affect-
ing gene transcription or transcription factors, the actual
binding targets in the nucleus are not known. We hypoth-
esized that AnkA would bind to DNA or DNA–protein
targets of granulocyte nuclei, and tested this by analysing
these binding properties in vitro. The results of these
analyses provide strong evidence of a DNA binding role
for AnkA, with a potential contribution by protein, and lead
to further speculation about the functional nature of
AnkA–host cell DNA interactions.