Starvation and rapamycin differentially regulate host
cell lysosome exocytosis and invasion by
Trypanosoma cruzi metacyclic forms
Rafael Miyazawa Martins, Renan Melatto Alves,
Silene Macedo and Nobuko Yoshida*
Departamento de Microbiologia, Imunologia e
Parasitologia, Universidade Federal de São Paulo,
R. Botucatu, 862, 6° andar, 04023-062 São Paulo,
Brasil.
Summary
The molecular mechanisms of host cell invasion
by T. cruzi metacyclic trypomastigotes (MT), the
developmental forms that initiate infection in the
mammalian host, are only partially understood.
Here we aimed at further identifying the target
cell components involved in signalling cascades
leading to MT internalization, and demonstrate for
the first time the participation of mammalian target
of rapamycin (mTOR). Treatment of human epithe-
lial HeLa cells with mTOR inhibitor rapamycin
reduced lysosomal exocytosis and MT invasion.
Downregulation of phosphatidylinositol 3-kinase
and protein kinase C also impaired exocytosis and
MT internalization. The recombinant protein based
on gp82, the MT surface molecule that mediates
cell adhesion/invasion, induced exocytosis in
HeLa cells. Such an effect has not previously been
attributed to any T. cruzi surface molecule. Rapa-
mycin treatment diminished gp82 binding as well.
Cell invasion assays under conditions that pro-
moted lysosome exocytosis, such as 1 h incuba-
tion in starvation medium PBS
++
, increased MT
invasion, whereas pre-starvation of cells for 1–2 h
had an opposite effect. In contrast to MT, invasion
of tissue culture trypomastigotes (TCT) increased
upon host cell pre-starvation or treatment with
rapamycin, a novel finding that discloses quite dis-
tinctive features of the two infective forms in a key
process for infection.
Introduction
The virulence of many bacteria and protozoan parasites
that are pathogenic to humans is critically dependent on
their ability to invade host cells. By exploiting host cell
components or even subverting cellular functions, many
of these microorganisms accomplish their internalization.
Re-arrangement of target cell actin cytoskeleton, for
instance, constitutes a strategy used by diverse entero-
pathogenic bacteria, such as Shigella, Salmonella. List-
eria and Yersisnia. To induce host cell actin remodelling,
Shigella and Salmonella use a specialized protein secre-
tion system to inject into target cells virulence factors that
trigger a variety of signalling pathways, whereas Listeria
and Yersinia engage transmembrane cell adhesion pro-
teins (Stebbins and Galán, 2001; Cossart and Sansonetti,
2004). Invasion of mammalian cells by the protozoan
parasite Trypanosoma cruzi requires the activation of sig-
nalling cascades that lead to intracellular Ca
2+
mobiliza-
tion (Docampo and Moreno, 1996; Burleigh and Andrews,
1998; Yoshida, 2006) and disruption of actin cytoskeleton
(Rodriguez et al., 1995; Cortez et al., 2006).
Trypanosoma cruzi causes Chagas’ disease, which
was predominantly a neglected disease of poor, rural and
forgotten populations in Latin America, but is now spread-
ing to non-endemic countries, posing a new worldwide
challenge (Coura and Viñas, 2010; Schmunis and Yadon,
2010). In endemic regions, oral transmission of T. cruzi
has been responsible for outbreaks of acute cases of
Chagas’ disease (Coura et al., 2002; Coura, 2006; Bastos
et al., 2010; Noya et al., 2010), with the involvement of
metacyclic forms from insect vectors. In natural infections,
metacyclic trypomastigotes (MT) are the parasite forms
that first contact the host cells. They are internalized in a
membrane-bounded vacuole, then escape to the cyto-
plasm and differentiate into the replicative amastigote
forms, which later transform into trypomastigotes that are
released in the circulation upon host cell rupture and
disseminate to diverse organs and tissues. Studies using
MT generated in liquid media and tissue culture-derived
trypomastigotes (TCT) as counterparts of insect-borne
and bloodstream parasites, respectively, have revealed
some common features of the mammalian cell invasion
(Burleigh and Andrews, 1998; Yoshida, 2006). However,
Received 22 November, 2010; revised 24 February, 2011; accepted
25 February, 2011. *For correspondence. E-mail nyoshida@
unifesp.br; Tel. (+55) 11 5549 5159; Fax (+55) 11 55711095.
Cellular Microbiology (2011) 13(7), 943–954 doi:10.1111/j.1462-5822.2011.01590.x
First published online 24 Apr 2011
© 2011 Blackwell Publishing Ltd
cellular microbiology