Gluthatione and Immunity

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ch \af0 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 \par }\pard \plain \qj \li0 \ri0 \sb174 \lisb0 \sa174 \lisa0 \widctl
par \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \
af0 \loch \af0 {\*\bkmkstart abstract0}{\*\bkmkend abstract0}{\b1 \fs31 \kerning
2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Abstract}{\b
1 \fs31 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2
052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \ri
n0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article
1.front1.article-meta1.abstract1.p1}{\*\bkmkend article1.front1.article-meta1.ab
stract1.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052
\langfenp2052 Glutathione (GSH), a tripeptide antioxidant, is essential for cell
ular homeostasis and plays a vital role in diverse cellular functions. Individua
ls who are infected with Human immuno deficiency virus (HIV) are known to be sus
ceptible to }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langf
e2052 \langfenp2052 Mycobacterium tuberculosis}{\fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 (}{\i1 \fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb)}{\fs21 \kerni
ng2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infection
. We report that by enhancing GSH levels, T-cells are able to inhibit the growth
of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \lang
fe2052 \langfenp2052 inside macrophages. In addition, those GSH-replenished T c
ell cultures produced increased levels of Interleukin-2 (IL-2), Interleukin-12 (
IL-12), and Interferon-gamma (IFN-\uc1 \u947 ?), cytokines, which are known to b
e crucial for the control of intracellular pathogens. Our study reveals that T l
ymphocytes that are derived from HIV infected individuals are deficient in GSH,
and that this deficiency correlates with decreased levels of Th1 cytokines and e
nhanced growth of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1
\langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0
\cgrid1 \langfe2052 \langfenp2052 inside human macrophages.}{\fs21 \kerning2 \
dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \pl
ain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs21 \kerning2 \dbch \af2 \hich \a
f0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Citation:}{\fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Guerra C, Morris D,
Sipin A, Kung S, Franklin M, Gray D, et al. (2011) Glutathione and Adaptive Immu
ne Responses against }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgri
d1 \langfe2052 \langfenp2052 Mycobacterium tuberculosis}{\fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Infection in Healthy
and HIV Infected Individuals. PLoS ONE 6(12): e28378. doi:10.1371/journal.pone.
0028378}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \la
ngfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \l
in0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Editor:
}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2
052 Shabaana Khader, University of Pittsburgh, United States of America}{\fs21 \
kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par
}\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0
\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs21 \kerning2 \dbch \af
2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Received:}{\fs21 \kerni
ng2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 June 28, 2
011; }{\b1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \
langfenp2052 Accepted:}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1
\langfe2052 \langfenp2052 November 7, 2011; }{\b1 \fs21 \kerning2 \dbch \af2 \hi
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Published:}{\fs21 \kerning2
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 December 2, 2
011}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfe
np2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0
\rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs21 \kernin
g2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Copyright:}
52 \uc1 \u169 ? 2011 Guerra et al. This is an open-access article distributed u
nder the terms of the Creative Commons Attribution License, which permits unrest
ricted use, distribution, and reproduction in any medium, provided the original
author and source are credited.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0
\cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \
aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \
loch \af0 {\b1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe20
52 \langfenp2052 Funding:}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgri
d1 \langfe2052 \langfenp2052 The study was supported by start-up funds from West
ern University of Health Sciences and Potts Memorial Foundation. The funders had
no role in study design, data collection and analysis, decision to publish, or
preparation of the manuscript.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0
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h \f0 \cgrid1 \langfe2052 \langfenp2052 Although, one of the co-authors of this
manuscript is a Managing Partner for Your Energy Systems (a small biotech compa
ny), this does not alter the authors' adherence to all the PLoS ONE policies on
sharing data and materials.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cg
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ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 HIV is the cause
of acquired immunodeficiency syndrome (AIDS). Both HIV-1 and HIV-2 cause AIDS, b
ut HIV-1 is found worldwide, whereas HIV-2 is found primarily in West Africa }{\
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fe2052 \langfenp2052 \cf13 \ul1 [3]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
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rgets for HIV infection and their role in AIDS pathogenesis are well documented
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angfe2052 \langfenp2052 \cf13 \ul1 [3]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0
\loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch
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\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052
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\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp20
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052 \langfenp2052 \cf13 \ul1 [5]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \
hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . HIV preferentially infec
ts and kills CD4+ T lymphocytes, resulting in the loss of cell-mediated immunity
and a high probability that the host will develop opportunistic infections incl
uding tuberculosis (TB).}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid
1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalph
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\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp205
2 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \la
ngfenp2052 is endemic in every part of the world. This organism accounts for ne
arly 1.7 million deaths annually, making it the leading bacterial cause of death
worldwide }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0
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\dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{
2 . Once thought to be controlled, the incidence of TB is rising in many areas c
aused in part by the emergence of drug-resistant strains and the HIV epidemic. F
urthermore, nearly one-third of the world is latently infected with }{\i1 \fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. t
b}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp
2052 , making eradication of the organism difficult }{\field \fldedit {\*\fldins
fe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/arti
cle?id=10.1371/journal.pone.0028378#pone.0028378-WHO1"}}{\*\fldrslt {\fs21 \kern
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np2052 \cf13 \ul1 [1]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid
1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \
loch \f0 \cgrid1 \langfe2052 \langfenp2052 .}{\fs21 \kerning2 \dbch \af2 \hich \
af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0
\widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2
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.body1.sec1.p3}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2
052 \langfenp2052 Although TB remains the leading cause of morbidity and mortali
ty due to any one infectious agent worldwide }{\field \fldedit {\*\fldinst {\fs2
1 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \
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\langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=
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52 \cf13 \ul1 [6]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \l
\f0 \cgrid1 \langfe2052 \langfenp2052 our understanding of its immunopathogene
sis is still incomplete.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid
f0 {\*\bkmkstart article1.body1.sec1.p4}{\*\bkmkend article1.body1.sec1.p4}{\i1
2 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \la
ngfenp2052 infection begins as primary TB with the deposition of bacilli in the
alveoli, which are phagocytosed by alveolar macrophages }{\field \fldedit {\*\f
ldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \la
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\langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone
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s21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052
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2 \langfenp2052 \cf13 \ul1 [7]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \a
f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hi
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Active TB is characterized
by a profound and prolonged suppression of }{\i1 \fs21 \kerning2 \dbch \af2 \hi
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 -specific T cell re
sponses, as evidenced by decreased production of the cytokines IL-2 and IFN-\uc1

\u947 ? }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \l
och \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \a
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0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [8]}}}{\fs21 \kerning2
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\langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
ch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [11]}}}{\fs21 \kerning2 \dbc
h \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs2
1 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ,
urnals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-Meh
ra1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \lan
f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [12]}}}{\fs21 \kerning2 \dbch \a
f2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \k
erning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Over
production of immunosuppressive cytokines [IL-10 and transforming growth factor
(TGF)- \uc1 \u946 ?] by mononuclear phagocytes has been implicated in decreased
T cell function during TB }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \
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\ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/journal.pon
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{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2
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loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \
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2 \langfenp2052 -}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich
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\af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2
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\kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf
13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 .}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe20
52 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustri
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\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 GSH plays a m
ajor role in the maintenance of the cellular redox state. GSH scavenges peroxide
species which can be harmful to the cells. GSH also plays a role in the normal
function of the immune system }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \db
ch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs
cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/journal
.pone.0028378#pone.0028378-Buhl1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [3]
}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfe
np2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \lan
gfe2052 \langfenp2052 . Of particular interest, is the ability of GSH to enhance
the activation of lymphocytes which play a major role in the pathology of HIV i
nfection. GSH has been demonstrated to be depleted in HIV positive individuals }
{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \
cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \
af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://jou
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\cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [3]}}}{\fs21 \kerning2 \dbch \af2
ing2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Low GSH
levels have been shown to result in the activation of nuclear factor \uc1 \u954
?B (NF\uc1 \u954 ?B), which is necessary for active transcription of the HIV pr
ovirus }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loc
h \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2
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tp://journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028
378-Staal1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgri
d1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [4]}}}{\fs21 \kerning2 \d
bch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\f
s21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052
, }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hic
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://
journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-S
taal2"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \l
\f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [5]}}}{\fs21 \kerning2 \dbch \
kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 , }{\
als.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-Duh1"}
grid1 \langfe2052 \langfenp2052 \cf13 \ul1 [16]}}}{\fs21 \kerning2 \dbch \af2 \h
ich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kernin
g2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Depleted
GSH levels have also been shown to play a role in the apoptosis of CD4}{\fs21 \k
erning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \super
1 +}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfe
np2052 T cells. As depletion of these T cells is the major pathology of the HIV
virus, replenishment of GSH represents an exciting prospect for treatment as a
supplement to highly active antiretroviral therapy (HAART) }{\field \fldedit {\*
\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \
langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid
1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/ploso
ne/article?id=10.1371/journal.pone.0028378#pone.0028378-Suthanthiran1"}}{\*\fldr
slt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langf
enp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \la
ngfe2052 \langfenp2052 \cf13 \ul1 [17]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 .}{\fs21 \kerning2 \
dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \pl
ain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec1.p6}{\*
\bkmkend article1.body1.sec1.p6}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0
\cgrid1 \langfe2052 \langfenp2052 Our earlier studies indicated that GSH facili
tates the control of growth of intracellular }{\i1 \fs21 \kerning2 \dbch \af2 \h
ich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbc
h \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 in both murine an
d human macrophages and has direct antimycobacterial activity }{\field \fldedit
{\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe205
2 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cg
rid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/pl
osone/article?id=10.1371/journal.pone.0028378#pone.0028378-Venketaraman1"}}{\*\f
ldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \la
\langfe2052 \langfenp2052 \cf13 \ul1 [18]}}}{\fs21 \kerning2 \dbch \af2 \hich \a
ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 -}{\field \fldedi
t {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2
052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \
cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/
plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-Dayaram1"}}{\*\fldr
ngfe2052 \langfenp2052 \cf13 \ul1 [20]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Furthermore, our r
ecent studies indicate that GSH in combination with IL-2 and IL-12 augments natu
ral killer (NK) cell functions to control }{\i1 \fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infection }{\field \
s.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-Millman1"}}{\
*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052
\langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgri
d1 \langfe2052 \langfenp2052 \cf13 \ul1 [21]}}}{\fs21 \kerning2 \dbch \af2 \hich
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 .}{\fs21 \kern
ing2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pa
rd \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs
21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec1.
p7}{\*\bkmkend article1.body1.sec1.p7}{\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
ch \f0 \cgrid1 \langfe2052 \langfenp2052 The ability of GSH to augment the activ
ity of NK cells in the control of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \l
och \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hic
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infection as indicated by ou
r previous studies }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hi
2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 H
YPERLINK "http://journals.plos.org/plosone/article?id=10.1371/journal.pone.00283
78#pone.0028378-Venketaraman1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \a
ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [18]}}
}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp
2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langf
e2052 \langfenp2052 -}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \
8378#pone.0028378-Dayaram1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [20]}}}{\
fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp205
52 \langfenp2052 led us to hypothesize that GSH will also augment the activity
of T-cells resulting in the control of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \a
f0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infection inside macrop
hages. Further, we believe that the decreased intracellular growth of }{\i1 \fs2
1 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M.
tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfe
np2052 will be accompanied by increased production of Th1 cytokines that are es
sential for the control of intracellular pathogens. We tested our hypothesis by
performing extensive }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgri
d1 \langfe2052 \langfenp2052 in vitro}{\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
ch \f0 \cgrid1 \langfe2052 \langfenp2052 studies using monocytes and T cells th
at are isolated from blood of healthy subjects and individuals with HIV infectio
n. Our results indicate that individuals with HIV infection have significantly l
ower levels of GSH in their T cells in comparison to healthy subjects. The decre
ase in the GSH levels correlated with increased growth of }{\i1 \fs21 \kerning2
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \
kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 insi
de human macrophages and reduced levels of Th1 cytokines in both plasma and cell
free supernatants derived from monocyte-T cell co-cultures. Our results signify
the importance of GSH in augmenting the functions of T lymphocytes to limit the
growth of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe
2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid
1 \langfe2052 \langfenp2052 inside monocytes and macrophages.}{\fs21 \kerning2
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \p
lain \qj \li0 \ri0 \sb174 \lisb0 \sa174 \lisa0 \widctlpar \aspalpha \adjustright
\lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmksta
rt s2}{\*\bkmkend s2}{\b1 \fs31 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid
1 \langfe2052 \langfenp2052 Results}{\b1 \fs31 \kerning2 \dbch \af2 \hich \af0 \
loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \sb2
10 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs
21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch \af2 \h
ich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Assay of GSH levels in T ce
lls from healthy and HIV-infected subjects}{\b1 \fs24 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \r
i0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \a
f2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec2.sec1.p1}{\*\bkmkend a
rticle1.body1.sec2.sec1.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgr
id1 \langfe2052 \langfenp2052 GSH levels were determined in freshly isolated T c
ells from thirteen healthy subjects and thirteen individuals with HIV infection.
We observed that GSH concentrations were significantly lower in T cells isolate
d from individuals with HIV infection compared to T cells from healthy subjects
(}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://j
ournals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g0
01"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \lang
fe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f
0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 1a}}}{\fs21 \kerning2 \dbc
1 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ).
The decreased levels of intracellular GSH in T cells from HIV-infected individu
als correlated with increased levels of TNF-\uc1 \u945 ? and free radicals. Decr
eased GSH levels in T cells of HIV positive individuals are likely to impair the
adaptive immune responses against }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \
loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hi
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infection. NAC-treatment of
T cells from individuals with HIV infection resulted in significant increase in
the levels of GSH (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \h
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52 \langfenp2052 ). Treatment of T cells from HIV positive individuals with BSO
resulted in significant decrease in the levels of GSH (}{\field \fldedit {\*\fld
inst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \lang
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\ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lan
gfenp2052 \cf13 \ul1 Figure 1b}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \a
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ).}{\fs21 \kerning2 \dbch \a
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2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj
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052 \langfenp2052 \cf13 \ul1 TIFF}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
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0 \ls1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 }{\field \fldedit {\
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052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustr
ight \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Figu
re 1. Assay of GSH levels in T cells isolated from healthy and HIV positive indi
viduals.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \
lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart
article1.body1.sec2.sec1.fig1.caption1.p1}{\*\bkmkend article1.body1.sec2.sec1.
fig1.caption1.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langf
e2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjus
tright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fs2
1 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 In
tracellular levels of GSH in freshly isolated T cells from healthy volunteers an
d HIV-infected individuals was determined by spectrophotometry, using an assay k
it from Calbiochem (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \h
378#pone-0028378-g001"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 1a}}}{\
52 \langfenp2052 ). T cells (2\uc1 \u215 ?10}{\fs21 \kerning2 \dbch \af2 \hich \
af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \super1 5}{\fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 /well) purified fro
m PBMCs using nylon wool columns were pelleted by centrifugation and an equal vo
lume of ice cold 5% MPA was added to the pellet. Supernatants were collected aft
er centrifugation and analyzed for total GSH using an assay kit from Calbiochem,
as per manufacturer\uc1 \u8217 ?s instructions. Total GSH in the samples were n
ormalized with protein. Proteins in the samples were estimated by Bradford\uc1 \
u8217 ?s method using Bio-Rad reagent. We also tested the effects of NAC and BSO
in increasing and decreasing the intracellular levels of GSH, respectively in T
cells isolated from both healthy subjects and individuals with HIV infection (}
rnals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g001
"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe
\cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 1b}}}{\fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). T
cells (2\uc1 \u215 ?10}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1
\langfe2052 \langfenp2052 \super1 5}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loc
h \f0 \cgrid1 \langfe2052 \langfenp2052 /well) isolated from healthy and HIV pos
itive subjects were treated as follows: mock treatment, treatment with NAC (10 m
M) and treatment with BSO (500 \uc1 \u181 ?M). Following overnight incubation, T
cells were pelleted and used for GSH measurement as per manufacturer\uc1 \u8217
?s instructions. Results shown in }{\field \fldedit {\*\fldinst {\fs21 \kerning
2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1
052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/jo
urnal.pone.0028378#pone-0028378-g001"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \
Figure 1b}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe20
grid1 \langfe2052 \langfenp2052 are averages from experiments performed using T
cells isolated from three healthy individuals and eight individuals with HIV in
fection. * represent significant difference in the levels of GSH between untreat
ed T cells derived from healthy subjects versus untreated T cells derived from i
ndividuals with HIV infection. **denotes significant difference in the levels of
GSH between untreated T cells versus NAC-treated T cells from HIV positive subj
ects. *** denotes statistically significant difference in the levels of GSH betw
een NAC-treated T cells and BSO-treated T cells from individuals with HIV infect
ion.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langf
enp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0
\rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\field \fldedit
rid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://dx.doi.org/10.1371/j
ournal.pone.0028378.g001"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \l
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 http://dx.d
oi.org/10.1371/journal.pone.0028378.g001}}}{\fs21 \kerning2 \dbch \af2 \hich \af
0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbc
\qj \li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \li
n0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \ker
ning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Assay of
free radicals and TNF-\uc1 \u945 ? in plasma samples}{\b1 \fs24 \kerning2 \dbch
\qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kernin
g2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec2.sec2.p1}{\
*\bkmkend article1.body1.sec2.sec2.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
ch \f0 \cgrid1 \langfe2052 \langfenp2052 MDA is a naturally formed byproduct of
the interaction between free radicals and lipid molecules which occurs in the bo
dy. The interaction between free radicals and lipid molecules produces lipid per
oxides which are unstable and decay to MDA and other products. Concentrations of
MDA are indicative of the level of cellular damage due to free radicals and the
refore are also indicative of levels of free radicals. Measurement of MDA concen
trations in the plasma of thirteen healthy and thirteen HIV infected individuals
revealed a significant increase in the amounts of MDA present in the plasma of
HIV infected individuals over those found in healthy individuals (}{\field \flde
dit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langf
\cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.or
g/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g002"}}{\*\fldrsl
fe2052 \langfenp2052 \cf13 \ul1 Figure 2a}}}{\fs21 \kerning2 \dbch \af2 \hich \a
ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). We observed an
eight-fold increase in the levels of TNF-\uc1 \u945 ? in the plasma samples of
HIV infected individuals compared to healthy subjects (}{\field \fldedit {\*\fld

rticle?id=10.1371/journal.pone.0028378#pone-0028378-g002"}}{\*\fldrslt {\fs21 \k
gfenp2052 \cf13 \ul1 Figure 2b}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \a
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). The difference in plasma
levels of TNF-\uc1 \u945 ? did not reach statistical significance due to a high
variance in the TNF-\uc1 \u945 ? concentrations measured in HIV infected individ
uals.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lang
fenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin
0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart po
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h \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2
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FF}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \lang
1 \langfe2052 \langfenp2052 }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbc
1 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \c
f13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article/figure/image?downlo
ad&size=original&id=info:doi/10.1371/journal.pone.0028378.g002"}}{\*\fldrslt {\f
2 \langfenp2052 \cf13 \ul1 original image () }}}{\fs21 \kerning2 \dbch \af2 \hic
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \
plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \
kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fs21 \kerning2 \dbch \af2 \hich \af0
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 Figure 2. Assay of free radicals an
d TNF-\uc1 \u945 ? in plasma samples from healthy and HIV positive individuals.}
52 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin
0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1
.body1.sec2.sec2.fig1.caption1.p1}{\*\bkmkend article1.body1.sec2.sec2.fig1.capt
ion1.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \la
ngfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \l
in0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fs21 \kernin
g2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Plasma samp
les separated from blood of healthy volunteers and HIV-infected individuals were
used for measurement of free radicals (}{\field \fldedit {\*\fldinst {\fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13
\ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lang
fenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.13
71/journal.pone.0028378#pone-0028378-g002"}}{\*\fldrslt {\fs21 \kerning2 \dbch \
\ul1 Figure 2a}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \lan
f0 \cgrid1 \langfe2052 \langfenp2052 ) and TNF-\uc1 \u945 ? (}{\field \fldedit {
sone/article?id=10.1371/journal.pone.0028378#pone-0028378-g002"}}{\*\fldrslt {\f
2 \langfenp2052 \cf13 \ul1 Figure 2b}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \l
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). Free radical levels
in plasma samples derived from healthy subjects and individuals with HIV infect
ion was determined by measuring the levels of MDA using a colorimetric assay kit
from Cayman. Levels of TNF-\uc1 \u945 ? in the plasma samples were determined b
y ELISA using assay kits procured from eBioscience. Results in }{\field \fldedit
grid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/p
losone/article?id=10.1371/journal.pone.0028378#pone-0028378-g002"}}{\*\fldrslt {
052 \langfenp2052 \cf13 \ul1 Figure 2}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 are averages of data
collected from thirteen healthy subjects and thirteen individuals with HIV infe
ction.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lan
gfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \li
n0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\field \fldedi
cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://dx.doi.org/10.1371
/journal.pone.0028378.g002"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 http://dx
.doi.org/10.1371/journal.pone.0028378.g002}}}{\fs21 \kerning2 \dbch \af2 \hich \
af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \d
bch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \pla
in \qj \li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \
lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \k
erning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Determ
ination of intracellular viability of }{\b1 \i1 \fs24 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\b1 \fs24 \kerning2 \db
ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 in monocyte-T ce
ll co-cultures from healthy and HIV positive individuals}{\b1 \fs24 \kerning2 \d
in \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \ker
ning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec2.sec3.p1
}{\*\bkmkend article1.body1.sec2.sec3.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 We then tested the intracellular sur
vival of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe20
52 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1
\langfe2052 \langfenp2052 in co-cultures of monocytes and T cells that were iso
lated from healthy subjects and individuals with HIV infection. Results from the
six healthy subjects revealed a two-fold increase in the growth of }{\i1 \fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. t
b}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp
2052 inside monocytes cultured both in the presence and absence of T cells (}{\
als.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g003"}
grid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3a}}}{\fs21 \kerning2 \dbch \a
erning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). Inc
ubation with NAC (10 mM)-treated T cells resulted in stasis in the growth of }{\
i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp
2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052
\langfenp2052 inside monocytes (}{\field \fldedit {\*\fldinst {\fs21 \kerning2
2 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/jour
nal.pone.0028378#pone-0028378-g003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hi
2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 F
igure 3a}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052
id1 \langfe2052 \langfenp2052 ). Incubation with BSO-treated T cells resulted in
abrogation in the growth inhibition and a six-fold growth of }{\i1 \fs21 \kerni
ng2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs
21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052
inside monocytes confirming that GSH enhances the functions of T lymphocytes to
control }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe205
2 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \
langfe2052 \langfenp2052 infection inside monocytes (}{\field \fldedit {\*\fldi
nst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langf
ngfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/ar
ticle?id=10.1371/journal.pone.0028378#pone-0028378-g003"}}{\*\fldrslt {\fs21 \ke
fenp2052 \cf13 \ul1 Figure 3a}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). Whereas in HIV positive in
dividuals, we observed a six-fold increase in the growth of }{\i1 \fs21 \kerning
2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 in
side monocytes cultured in the absence of T cells (}{\field \fldedit {\*\fldinst
{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp
e2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/artic
le?id=10.1371/journal.pone.0028378#pone-0028378-g003"}}{\*\fldrslt {\fs21 \kerni
ng2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul
1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfen
p2052 \cf13 \ul1 Figure 3b}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \
af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ) and five-fold increase in the
growth of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2
052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1
\langfe2052 \langfenp2052 in monocytes cultured in the presence of mock-treate
d T cells (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0
"http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone0028378-g003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cg
0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3b}}}{\fs21 \ker
enp2052 ). These results indicate that monocytes and T cells from HIV positive i
ndividuals are unable to restrict the growth of }{\i1 \fs21 \kerning2 \dbch \af2
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \
dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 to the same ex
tent as healthy individuals. Treatment of T cells from HIV positive subjects wit
h 10 mM NAC and co-incubation with infected monocytes resulted in reduction in t
he fold increase in growth of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \a
f0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 inside monocytes compared to oth
er treatment conditions. However, NAC (10 mM)-treatment of T cells did not resul
t in complete stasis in the growth of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af
0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 as seen in healthy subje
cts (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http
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8-g003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \
langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loc
h \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3b}}}{\fs21 \kerning2
2 ). We also observed a poor uptake of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \a
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 by monocytes derived fr
om individuals with HIV infection compared to healthy individuals as evidence by
lower colony counts recovered from the co-cultures of monocytes and T cells (}{
\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \c
f0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://jour
nals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g003"
}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2
cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3b}}}{\fs21 \kerning2 \dbch \
kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). Th
is reduced uptake of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgri
d1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\f0 \cgrid1 \langfe2052 \langfenp2052 could possibly be due to impaired phagocy
tic capacity of monocytes that are derived from HIV positive individuals.}{\fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \pa
r }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \ita
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ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \af0 {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgri
d1 \langfe2052 \langfenp2052 Figure 3. Intracellular survival of H37Rv inside T
cell-monocyte co-cultures.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgr
id1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspal
pha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch
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r \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af
0 \loch \af0 {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe205
2 \langfenp2052 We determined the intracellular survival of H37Rv inside T cellmonocyte co-cultures from healthy subjects (}{\field \fldedit {\*\fldinst {\fs21
\kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \c
f13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \
langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=1
0.1371/journal.pone.0028378#pone-0028378-g003"}}{\*\fldrslt {\fs21 \kerning2 \db
cf13 \ul1 Figure 3a}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1
ch \f0 \cgrid1 \langfe2052 \langfenp2052 ) and individuals with HIV infection (}
rnals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g003
"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). H
uman monocytes were infected with the processed virulent laboratory strain of }{
\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfen
p2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052
\langfenp2052 , H37Rv at a multiplicity of infection of 10\uc1 \u8758 ?1. T cel
ls were purified using nylon wool column. T cells were treated as follows: a) no
additives b) NAC (5 mM) c) NAC (10 mM) d) NAC (20 mM) and e) BSO (500 \uc1 \u18
1 ?M) for 24 h. T cells were washed, resuspended in fresh media and then added t
o the infected monocytes. Infected monocytes-T cell co-cultures were terminated
at 1 hour and 5 days post-infection to determine the intracellular survival of H
37Rv inside human monocytes. Monocyte lysates were plated on 7H11 medium enriche
d with ADC to estimate the growth or killing of H37Rv. Results shown in }{\field
\fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1
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los.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g003"}}{\*\
\langfe2052 \langfenp2052 \cf13 \ul1 Figure 3a}}}{\fs21 \kerning2 \dbch \af2 \h
g2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 are averag
es from five different experiments performed in triplicate. Results shown in }{\
grid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3b}}}{\fs21 \kerning2 \dbch \a
erning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 are a
verages from seven different experiments performed in triplicate.}{\fs21 \kernin
g2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard
\plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21
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.g003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \l
\f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 http://dx.doi.org/10.1371/jour
nal.pone.0028378.g003}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid
21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch \af2 \h
ich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Assay of IL-12, IL-2 and IF
N-\uc1 \u947 ? in supernatants derived from T cell-monocyte co-cultures}{\b1 \fs
tap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.bod
y1.sec2.sec4.p1}{\*\bkmkend article1.body1.sec2.sec4.p1}{\fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 To determine whether
the growth inhibition of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \
cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \l
och \f0 \cgrid1 \langfe2052 \langfenp2052 in monocyte-T cell co-cultures is acc
ompanied by increased production of IL-12, IL-2 and IFN-\uc1 \u947 ?, we quantif
ied the levels of these cytokines in the cell free-supernatants from co-cultures
of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \lang
fe2052 \langfenp2052 -infected monocytes and T cells (from healthy subjects and
individuals with HIV infection) by ELISA (}{\field \fldedit {\*\fldinst {\fs21 \
kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf1
3 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \la
ngfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.
1371/journal.pone.0028378#pone-0028378-g004"}}{\*\fldrslt {\fs21 \kerning2 \dbch
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ngfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\f0 \cgrid1 \langfe2052 \langfenp2052 ). We observed an increase in the producti
on of IL-12, IL-2 and IFN-\uc1 \u947 ? in monocyte+NAC-treated T cell co-culture
s derived from healthy subjects (}{\field \fldedit {\*\fldinst {\fs21 \kerning2
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GSH in T cells favors the synthesis of Th-1 cytokines leading to the control of
}{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lang
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52 \langfenp2052 infection. We also quantified the levels of IL-12, IL-2 and IF

N-\uc1 \u947 ? in monocyte-T cell co-cultures from individuals with HIV infectio
n (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \a
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\f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 4d and e}}}{\fs21 \kerni
p2052 ). We were unable to detect IL-12 in the culture supernatants. IL-2 and IF
N-\uc1 \u947 ? were detected in monocyte-T cell co-cultures however the levels o
f these cytokines were almost one log lower compared to the healthy individuals
(}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0
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0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 4d and e}}}{\fs21 \kerning
052 ). This signifies the consequences of HIV infection on T cell functions. Sin
ce mycobacteria are not internalized by T cells the downregulation in the produc
tion of IFN-\uc1 \u947 ? and IL-2 by T cells is an HIV induced effect.}{\fs21 \k
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ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Figure 4. Assay of IL-12, IL
-2 and IFN-\uc1 \u947 ?in supernatants derived from co-cultures of T cells-}{\i1
52 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
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gfe2052 \langfenp2052 Levels of IL-12, IL-2 and IFN-\uc1 \u947 ? were assayed in
supernatants derived from co-cultures of T cells-}{\i1 \fs21 \kerning2 \dbch \a
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 -infected mon
ocytes from healthy individuals (}{\field \fldedit {\*\fldinst {\fs21 \kerning2
igure 4a, b and c}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \l
\f0 \cgrid1 \langfe2052 \langfenp2052 ) and individuals with HIV infection (}{\
grid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 4d and e}}}{\fs21 \kerning2 \d
) T cells were purified using nylon wool column. T cells were treated as follows
: 1) no additives 2) NAC (5 mM) 3) NAC (10 mM) 4) NAC (20 mM) and 5) BSO (500 \u
c1 \u181 ?M) for 24 h. Following incubation with stimulants, T cells were washed
, re-suspended in fresh RPMI containing human serum without any stimulants and t
hen added to the infected monocytes. Supernatants were collected from co-culture
s of H37Rv-infected monocytes-T cells at 5 days post-infection were filtered and
assayed for the levels of IL-12, IL-2 and IFN-\uc1 \u947 ? using assay kits fro
m eBioscience. Data in }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2
\hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 }{\fs21 \ker
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028378#pone-0028378-g004"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \l
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figures 4a,
b and c}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052
id1 \langfe2052 \langfenp2052 represent means\uc1 \u177 ?SE from four different
healthy individuals. Data in }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \db
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res 4d and e}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe
\cgrid1 \langfe2052 \langfenp2052 represent means\uc1 \u177 ?SE from six differ
ent individuals with HIV infection.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
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ngfe2052 \langfenp2052 Assay of IL-12, IL-2, IFN-\uc1 \u947 ? and IL-10 levels i
n plasma samples from healthy and HIV-infected subjects}{\b1 \fs24 \kerning2 \db
ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plai
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{\*\bkmkend article1.body1.sec2.sec5.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \
loch \f0 \cgrid1 \langfe2052 \langfenp2052 Measurement of Th-1 cytokines such as
IL-12, IL-2 and IFN-\uc1 \u947 ? in plasma samples from thirteen healthy and th
irteen HIV infected individuals revealed a significant reduction in the levels o
f these cytokines in HIV infected individuals when compared to healthy subjects.
We observed a statistically significant (i.e., 50%) decrease in the levels of I
L-12 in plasma samples from individuals with HIV infection compared to healthy s
ubjects (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \l
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-00
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p2052 ). Decreased levels of IL-12 will interfere with the differentiation of Th
0 to Th1 subset }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich
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pone.0028378-Clerici1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
angfenp2052 . We also observed a 20% decrease in the levels of IL-2 in plasma sa
mples derived from individuals with HIV infection compared to healthy subjects (
urnals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g00
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ).D
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s }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
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\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Fu
rthermore, we observed a significant decrease in the levels of IFN-\uc1 \u947 ?
in plasma samples from individuals with HIV infection compared to healthy subjec
ts (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \
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10.1371/journal.pone.0028378.g005"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hic
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 or
iginal image () }}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \la
\f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpa
r \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af
0 \loch \af0 {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe205
2 \langfenp2052 Figure 5. Assay of IL-12, IL-2, IFN-\uc1 \u947 ? and IL-10 in pl
asma samples from healthy and HIV positive individuals.}{\fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \q
j \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2
\dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec2.sec5.fig1.ca
ption1.p1}{\*\bkmkend article1.body1.sec2.sec5.fig1.caption1.p1}{\fs21 \kerning2
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 Plasma samples separated from blood
of healthy volunteers and HIV-infected individuals were used for measurement of
IL-12 (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "h
ttp://journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-002
8378-g005"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid
loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 5a}}}{\fs21 \kernin
2052 ), IL-2 (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \a
INK "http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#po
ne-0028378-g005"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 5b}}}{\fs21 \
ngfenp2052 ), IFN-\uc1 \u947 ? (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \
\cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/journ
al.pone.0028378#pone-0028378-g005"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hic
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Fi
gure 5c}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052
d1 \langfe2052 \langfenp2052 ) and IL-10 (}{\field \fldedit {\*\fldinst {\fs21 \
1371/journal.pone.0028378#pone-0028378-g005"}}{\*\fldrslt {\fs21 \kerning2 \dbch
13 \ul1 Figure 5d}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \l
\f0 \cgrid1 \langfe2052 \langfenp2052 ) by ELISA using assay kits from eBioscie
nce. Results in }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich
RLINK "http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#
pone-0028378-g005"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 5}}}{\fs21
angfenp2052 are averages from data collected from thirteen healthy subjects and
thirteen individuals with HIV infection.}{\fs21 \kerning2 \dbch \af2 \hich \af0
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \w
idctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \h
ich \af0 \loch \af0 {\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hi
YPERLINK "http://dx.doi.org/10.1371/journal.pone.0028378.g005"}}{\*\fldrslt {\fs
\langfenp2052 \cf13 \ul1 http://dx.doi.org/10.1371/journal.pone.0028378.g005}}}
2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \sb174 \lisb0 \sa174 \lisa0
\hich \af0 \loch \af0 {\*\bkmkstart s3}{\*\bkmkend s3}{\b1 \fs31 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Discussion}{\b1 \fs
tap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.bod
y1.sec3.p1}{\*\bkmkend article1.body1.sec3.p1}{\fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Blood monocytes and resident ma
crophages are important }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \c
grid1 \langfe2052 \langfenp2052 in vivo}{\fs21 \kerning2 \dbch \af2 \hich \af0 \
loch \f0 \cgrid1 \langfe2052 \langfenp2052 cell targets for HIV infection and t
heir role in AIDS pathogenesis is well documented }{\field \fldedit {\*\fldinst
2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/articl
e?id=10.1371/journal.pone.0028378#pone.0028378-Zhang1"}}{\*\fldrslt {\fs21 \kern
np2052 \cf13 \ul1 [2]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid
loch \f0 \cgrid1 \langfe2052 \langfenp2052 . These cells of innate immune defens
es usually survive HIV infection, serve as a major virus reservoir, and function
as immunoregulatory cells through secretion of several pro-inflammatory cytokin
es and chemokines in response to HIV infection, thereby recruiting and activatin
g CD4+ T cells which serve as new target cells for the virus }{\field \fldedit {
sone/article?id=10.1371/journal.pone.0028378#pone.0028378-Buhl1"}}{\*\fldrslt {\
52 \langfenp2052 \cf13 \ul1 [3]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \
ich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 -}{\field \fldedit {\*\fldi
ngfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/ar
ticle?id=10.1371/journal.pone.0028378#pone.0028378-Staal2"}}{\*\fldrslt {\fs21 \
ngfenp2052 \cf13 \ul1 [5]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \c
f0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 .}{\fs21 \kerning2 \dbch \af2 \hi
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0
\ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec3.p2}{\*\bkmkend arti
cle1.body1.sec3.p2}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \lan
gfe2052 \langfenp2052 GSH is the primary antioxidant compound employed by human
cells in the maintenance of the cellular redox balance. GSH is a tripeptide made
of glutamine, cysteine, and glycine and has been shown to be important for norm
al function of the immune system. In particular, we have previously demonstrated
that GSH is important for the control of intracellular }{\i1 \fs21 \kerning2 \d
bch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infect
ion by the cells of the immune system }{\field \fldedit {\*\fldinst {\fs21 \kern
np2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371
/journal.pone.0028378#pone.0028378-Venketaraman1"}}{\*\fldrslt {\fs21 \kerning2
\f0 \cgrid1 \langfe2052 \langfenp2052 -}{\field \fldedit {\*\fldinst {\fs21 \ke
fenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.13
71/journal.pone.0028378#pone.0028378-Millman1"}}{\*\fldrslt {\fs21 \kerning2 \db
cf13 \ul1 [21]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \lang
0 \cgrid1 \langfe2052 \langfenp2052 . Conditions of depleted GSH have been demon
strated to activate transcription factors that are necessary for active replicat
ion of the HIV within lymphocytes [3-5, 16 &17].}{\fs21 \kerning2 \dbch \af2 \hi
\af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec3.p3}{\*\bkmkend arti
cle1.body1.sec3.p3}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \lan
gfe2052 \langfenp2052 In this study, we observed that GSH levels are compromised
in T lymphocytes derived from individuals with HIV infection (}{\field \fldedit
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ) and this decrease c
orrelated with increased levels of TNF-\uc1 \u945 ? and free radicals in the pla
sma compared to healthy subjects (}{\field \fldedit {\*\fldinst {\fs21 \kerning2
52 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/jou
rnal.pone.0028378#pone-0028378-g002"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \h
Figure 2}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052
id1 \langfe2052 \langfenp2052 ).}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f
0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar
\loch \af0 {\*\bkmkstart article1.body1.sec3.p4}{\*\bkmkend article1.body1.sec3.
p4}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfen
p2052 TNF-\uc1 \u945 ?, a pro-inflammatory cytokine is considered to play a crit
ical role in the origin and progression of HIV infection }{\field \fldedit {\*\f
/article?id=10.1371/journal.pone.0028378#pone.0028378-Bahia1"}}{\*\fldrslt {\fs2
\langfenp2052 \cf13 \ul1 [23]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . The immuno-regulatory respo
nse of the host influences the pathogenesis of HIV-1 infection, triggering monoc
ytes, macrophages, and natural killer cells to produce TNF-\uc1 \u945 ? }{\field
\fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1
ch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.p
los.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-Alfano1"}}{
id1 \langfe2052 \langfenp2052 \cf13 \ul1 [24]}}}{\fs21 \kerning2 \dbch \af2 \hic
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . As a result
, there is a positive correlation between HIV-1 viremia and TNF-\uc1 \u945 ? lev
els in serum of HIV-1 infected patients. This relationship suggests that reducin
g TNF-\uc1 \u945 ? levels may also reduce occurrence of HIV-1 viremia. In excess
, TNF-\uc1 \u945 ? may cause severe inflammatory damage and toxicity, making con
trol of its production and secretion highly important. Regulating its release se
rves as a potential means of therapy for HIV-1. TNF-\uc1 \u945 ? can also induce
other pro-inflammatory cytokines such as IL-6 and IL-8, which aid in the upregu
lation of viral replication }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch
13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/journal.p
one.0028378#pone.0028378-FernandezOrtega1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \
\ul1 [25]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe20
grid1 \langfe2052 \langfenp2052 . Studies have also shown the ability of TNF-\uc
1 \u945 ? to stimulate production of anti-inflammatory cytokine IL-10, preventin
g further inflammation by causing TNF-\uc1 \u945 ? inhibition }{\field \fldedit
osone/article?id=10.1371/journal.pone.0028378#pone.0028378-Leghmari1"}}{\*\fldrs
lt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfe
gfe2052 \langfenp2052 \cf13 \ul1 [26]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . TNF-\uc1 \u945 ? is
secreted during the early phase of acute inflammatory diseases. Its pathogenic
role in HIV-1 infection involves activation of NF-\uc1 \u954 ?B, stimulating apo
ptosis of T lymphocytes. Tissue and plasma samples of hosts express high levels
of TNF-\uc1 \u945 ?, contributing to fever, anorexia, and other symptoms of HIV/
AIDS.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lang
fenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin
0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart ar
ticle1.body1.sec3.p5}{\*\bkmkend article1.body1.sec3.p5}{\fs21 \kerning2 \dbch \
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 It has been shown tha
t HIV infection results in increased production of free radicals and TNF-\uc1 \u
945 ? by macrophages }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \
8378#pone.0028378-Bahia1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \l
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [23]}}}{\fs
\langfenp2052 , }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich
#pone.0028378-Alfano1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
angfenp2052 . TNF-\uc1 \u945 ? stimulates the production of free radicals. Moreo
ver, enhanced levels of free radicals are likely to increase TNF-\uc1 \u945 ? in
various cells.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe
2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjust
right \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bk
mkstart article1.body1.sec3.p6}{\*\bkmkend article1.body1.sec3.p6}{\fs21 \kernin
g2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Chronic oxi
dative stress is often associated with HIV infection and research indicates a be
nefit for increased antioxidant vitamins and supplements in reduction of DNA bas
e damage, which in turn can slow progression of infection }{\field \fldedit {\*\
fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \l
\langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/ploson
e/article?id=10.1371/journal.pone.0028378#pone.0028378-Jaruga1"}}{\*\fldrslt {\f
2 \langfenp2052 \cf13 \ul1 [27]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \
ich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . The progression of HIV is
correlated with a decreased immunity. One way in which this decreased immunity
progresses is by free radical overload of monocytes and granulocytes leads to de
ficiency of antioxidant mechanisms which, in turn, may lead to the loss of CD4 c
ells often seen in the progression of HIV }{\field \fldedit {\*\fldinst {\fs21 \
1371/journal.pone.0028378#pone.0028378-Dobmeyer1"}}{\*\fldrslt {\fs21 \kerning2
\f0 \cgrid1 \langfe2052 \langfenp2052 . The decreased immunity may also be rela
ted to the reactive oxygen species and free radical presence which is higher in
HIV infected patients. With HIV infection progression there is an increased prod
uction of reactive oxygen species which leads to the theory of free radical medi
ated apoptosis of lymphocytes which reduces the ability for immune response to p
rogressive HIV infections }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \
\ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/journal.pon
e.0028378#pone.0028378-Dobmeyer1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [28
]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langf
ngfe2052 \langfenp2052 . With regards to CD4 T cell counts the apoptosis of lymp
hocytes by free radicals leads to progression of immunodeficiency and makes for
a quicker transition from HIV infection to AIDS }{\field \fldedit {\*\fldinst {\
52 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?
id=10.1371/journal.pone.0028378#pone.0028378-Bautista1"}}{\*\fldrslt {\fs21 \ker
enp2052 \cf13 \ul1 [29]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgr
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Furthermore, there is a link to l
ipid peroxidation observed in patients with HIV or AIDS to a deficiency of antio
xidants which leads to free radical proliferation }{\field \fldedit {\*\fldinst

2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/articl
e?id=10.1371/journal.pone.0028378#pone.0028378-Favier1"}}{\*\fldrslt {\fs21 \ker
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 .}{\fs21 \kerning2 \dbch \af2 \hich
f2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec3.p7}{\*\bkmkend articl
e1.body1.sec3.p7}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langf
e2052 \langfenp2052 Results of the co-culture studies using monocytes and T cell
s from healthy individuals indicate a complete inhibition in the growth of }{\i1
52 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 when infected monocytes were co-incubated with 10 mM NAC-treated Tcells (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loc
tp://journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028
378-g003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1
och \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3a}}}{\fs21 \kerning
052 ). Furthermore, co-incubation of infected monocytes with BSO-treated T cells
demonstrated six-fold increase in the intracellular growth of }{\i1 \fs21 \kern
ing2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\f
(}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g
003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \lan
f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3a}}}{\fs21 \kerning2 \db
21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 )
. In contrast to healthy subjects, we observed a several-fold increase in the gr
owth of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe205
2 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \
langfe2052 \langfenp2052 in co-cultures of monocytes and T cells derived from i
ndividuals with HIV infection (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \d
\cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/journa
l.pone.0028378#pone-0028378-g003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Fig
ure 3b}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \
1 \langfe2052 \langfenp2052 ). Treatment of T cells derived from HIV positive in
dividuals with 10 mM NAC resulted in reduction in the fold increase in growth of
}{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lang
fenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2
052 \langfenp2052 inside monocytes compared to other treatment conditions but d
id not result in complete stasis in the growth of }{\i1 \fs21 \kerning2 \dbch \a
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2

\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 as seen in h
ealthy subjects (}{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich
#pone-0028378-g003"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \a
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 3b}}}{\fs2
\langfenp2052 ). This is the first study to demonstrate that treatment with a GS
H-enhancing agent increases the functional activity of human T-cells resulting i
n the control of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \
langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0
\cgrid1 \langfe2052 \langfenp2052 infection. Our results indicate that treatmen
t of T cells from healthy subjects with NAC leads to the increased production of
IL-12, IL-2 and IFN-\uc1 \u947 ? (}{\field \fldedit {\*\fldinst {\fs21 \kerning
052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/jo
urnal.pone.0028378#pone-0028378-g004"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \
Figure 4a}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe20
grid1 \langfe2052 \langfenp2052 ). These cytokines drive the Th1 response and ar
e crucially important for the control of intracellular pathogens like }{\i1 \fs2
tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfe
np2052 .}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \
lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart
article1.body1.sec3.p8}{\*\bkmkend article1.body1.sec3.p8}{\fs21 \kerning2 \dbc
h \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 In our study, we a
lso observed decreased levels of IL-12, IL-2 and IFN-\uc1 \u947 ? in plasma samp
les from individuals with HIV infection compared to healthy subjects (}{\field \
s.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g005"}}{\*\fl
langfe2052 \langfenp2052 \cf13 \ul1 Figure 5a, 5b and 5c}}}{\fs21 \kerning2 \dbc
1 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ).
Activated antigen presenting cells secrete IL-12 which causes Th cell different
iation into the Th1 subset of cells }{\field \fldedit {\*\fldinst {\fs21 \kernin
2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id=10.1371/j
ournal.pone.0028378#pone.0028378-Clerici1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \
\ul1 [22]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe20
grid1 \langfe2052 \langfenp2052 . These Th1 cells then secrete a characteristic
Th1 profile of cytokines consisting of IL-2 and IFN-\uc1 \u947 ?. IL-2 induces p
roliferation of na\uc1 \u239 ?ve Th cells (Th}{\fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \sub2 0}{\fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ), amplifying the Th
response. IFN-\uc1 \u947 ? induces further IL-12 production in activated antige
n presenting cells, amplifying the Th1 response, and suppressing any Th2 respons
e }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
journals.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-H
uygen1"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \
h \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 [10]}}}{\fs21 \kerning2 \dbch
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . I
FN-\uc1 \u947 ? also plays an important role in the activation of innate immune
cells such as macrophages to control intracellular infections }{\field \fldedit
osone/article?id=10.1371/journal.pone.0028378#pone.0028378-Huygen1"}}{\*\fldrslt
e2052 \langfenp2052 \cf13 \ul1 [10]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 , }{\field \fldedit {\*
1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/ploso
ne/article?id=10.1371/journal.pone.0028378#pone.0028378-Osakwe1"}}{\*\fldrslt {\
hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Our results are consiste
nt with other published reports that in individuals infected with HIV, the norma
l Th1 response to viral infection is shifted to a Th2 response }{\field \fldedit
losone/article?id=10.1371/journal.pone.0028378#pone.0028378-Osakwe1"}}{\*\fldrsl
fe2052 \langfenp2052 \cf13 \ul1 [31]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \l
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 , }{\field \fldedit {\
d1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plos
one/article?id=10.1371/journal.pone.0028378#pone.0028378-Klein1"}}{\*\fldrslt {\
hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 .}{\fs21 \kerning2 \dbch \
j \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2
\dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec3.p9}{\*\bkmke
nd article1.body1.sec3.p9}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgri
d1 \langfe2052 \langfenp2052 We also observed increased levels of IL-10 in plasm
a samples from individuals with HIV infection compared to healthy subjects (}{\f
ield \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgr
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journa

ls.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g005"}}
{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe205
rid1 \langfe2052 \langfenp2052 \cf13 \ul1 Figure 5d}}}{\fs21 \kerning2 \dbch \af
rning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). IL-1
0 inhibits T cell proliferation and IFN-\uc1 \u947 ? production. Elevated levels
of IL-10 in serum during advanced HIV infection may enhance immune suppression,
allowing for opportunistic infections.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \
loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \wid
ctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hic
h \af0 \loch \af0 {\*\bkmkstart article1.body1.sec3.p10}{\*\bkmkend article1.bod
y1.sec3.p10}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052
\langfenp2052 Programmed death-1 (PD-1) and IL-10 are both upregulated during H
IV infection. Blocking interactions between PD-1 and programmed death ligand-1 (
PD-L1) and between IL-10 and IL-10 receptor (IL-10R) results in viral clearance
and improves T cell function in animal models of chronic viral infections }{\fie
ld \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid
loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals
.plos.org/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-Said1"}}{
id1 \langfe2052 \langfenp2052 \cf13 \ul1 [33]}}}{\fs21 \kerning2 \dbch \af2 \hic
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Additionall
y, blockade of the IL-10 pathway augmented }{\i1 \fs21 \kerning2 \dbch \af2 \hic
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 in vitro}{\fs21 \kerning2 \db
ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 proliferative ca
pacity of HIV-specific CD4 and CD8 T cells in individuals with HIV. IL-10 blocka
de also increased cytokine secretion by HIV-specific CD4 T cells }{\field \flded
\cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org
/plosone/article?id=10.1371/journal.pone.0028378#pone.0028378-Brockman1"}}{\*\fl
langfe2052 \langfenp2052 \cf13 \ul1 [34]}}}{\fs21 \kerning2 \dbch \af2 \hich \af
h \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . Our results ther
efore confirm the previous findings of other investigators that the levels of IL
-12, IL-2 and IFN-\uc1 \u947 ? are decreased in the plasma samples of individual
s with HIV infection (who in turn have higher IL-10) }{\field \fldedit {\*\fldin
st {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \langfe
gfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/art
icle?id=10.1371/journal.pone.0028378#pone.0028378-Klein1"}}{\*\fldrslt {\fs21 \k
gfenp2052 \cf13 \ul1 [32]}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \c
f0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 -}{\field \fldedit {\*\fldinst {\
52 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?
id=10.1371/journal.pone.0028378#pone.0028378-Brockman1"}}{\*\fldrslt {\fs21 \ker
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 .}{\fs21 \kerning2 \dbch \af2 \hich
f2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec3.p11}{\*\bkmkend artic
le1.body1.sec3.p11}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \lan
gfe2052 \langfenp2052 Further, our results show that individuals infected with H
IV have significantly lower levels of GSH within their T cells in comparison to
healthy subjects and this decrease correlated with reduced production of Th1 cyt
okines and compromised control of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \l
och \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hic
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infection (}{\field \fldedit
af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 , }{\field \fldedit {
sone/article?id=10.1371/journal.pone.0028378#pone-0028378-g003"}}{\*\fldrslt {\f
2 \langfenp2052 \cf13 \ul1 3}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 , }{\field \fldedit {\*\fldins
fe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/arti
cle?id=10.1371/journal.pone.0028378#pone-0028378-g004"}}{\*\fldrslt {\fs21 \kern
np2052 \cf13 \ul1 4}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1
ch \f0 \cgrid1 \langfe2052 \langfenp2052 and }{\field \fldedit {\*\fldinst {\fs
\langfenp2052 \cf13 \ul1 HYPERLINK "http://journals.plos.org/plosone/article?id
=10.1371/journal.pone.0028378#pone-0028378-g005"}}{\*\fldrslt {\fs21 \kerning2 \
\cf13 \ul1 5}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langf
\cgrid1 \langfe2052 \langfenp2052 ).}{\fs21 \kerning2 \dbch \af2 \hich \af0 \lo
ch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widct
lpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \af0 {\*\bkmkstart article1.body1.sec3.p12}{\*\bkmkend article1.body1
.sec3.p12}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \
langfenp2052 To conclude, our results indicate that HIV enhances the synthesis o
f TNF-\uc1 \u945 ? and free radicals leading to decreased levels of GSH in T lym
phocytes. The decreased GSH levels cause impaired production of Th1 cytokines re
sulting in enhanced growth of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \a
f0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 inside macrophages (}{\field \fl

org/plosone/article?id=10.1371/journal.pone.0028378#pone-0028378-g006"}}{\*\fldr
ngfe2052 \langfenp2052 \cf13 \ul1 Figure 6}}}{\fs21 \kerning2 \dbch \af2 \hich \
bch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 ). Our findings
suggest the possibility of efficacy for supplemental GSH therapy in HIV-}{\i1 \f
M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lang
fenp2052 co-infected individuals. In our future studies, we will further charac
terize the effects of decreased GSH in altering the functional activity of vario
us T cell sub-populations.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgr
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fe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adju
stright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fi
eld \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgri
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s.plos.org/plosone/article/figure/image?download&size=original&id=info:doi/10.13
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h \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 TIFF}}}
ngfe2052 \langfenp2052 }{\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af
ul1 HYPERLINK "http://journals.plos.org/plosone/article/figure/image?download&si
ze=original&id=info:doi/10.1371/journal.pone.0028378.g006"}}{\*\fldrslt {\fs21 \
ngfenp2052 \cf13 \ul1 original image () }}}{\fs21 \kerning2 \dbch \af2 \hich \af
\qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerni
ng2 \dbch \af2 \hich \af0 \loch \af0 {\fs21 \kerning2 \dbch \af2 \hich \af0 \loc
h \f0 \cgrid1 \langfe2052 \langfenp2052 Figure 6. Model describing the mechanism
by which GSH-enhanced T cells control }{\i1 \fs21 \kerning2 \dbch \af2 \hich \a
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 infection in macrophage
s.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfen
p2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \
rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\field \fldedit {\
d1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://dx.doi.org/10.1371/jou
rnal.pone.0028378.g006"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loc
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 http://dx.doi
.org/10.1371/journal.pone.0028378.g006}}}{\fs21 \kerning2 \dbch \af2 \hich \af0
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \
qj \li0 \ri0 \sb174 \lisb0 \sa174 \lisa0 \widctlpar \aspalpha \adjustright \lin0
\rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart s4}
{\*\bkmkend s4}{\b1 \fs31 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \lan
gfe2052 \langfenp2052 Materials and Methods}{\b1 \fs31 \kerning2 \dbch \af2 \hic
ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \i
tap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Subjects}{\b1 \fs24
p0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1
.sec4.sec1.p1}{\*\bkmkend article1.body1.sec4.sec1.p1}{\fs21 \kerning2 \dbch \af
2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 A total of 26 volunteer
s (13 healthy subjects and 13 individuals with HIV infection) were recruited for
the study. Individuals with HIV infection were recruited from the Foothills AID
S project. Healthy subjects without HIV infection or a history of TB were recrui
ted from the university faculty and staff. All HIV-infected volunteers had been
diagnosed with HIV-1, were taking some form of anti-retroviral treatment (ART),
and had CD4+ T-cell counts between 271 and 1415 cells per mm}{\fs21 \kerning2 \d
bch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \super1 3}{\fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 . T
hirty five milli-liters of blood was drawn once from both healthy volunteers and
individuals with HIV infection after obtaining a signed informed consent. All o
ur studies were approved by both the Institutional Review Board and the Institut
ional Biosafety Committee of Western University of Health Sciences.}{\fs21 \kern
rd \plain \qj \li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjust
right \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \
Separation of blood components}{\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch
\f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlp
ar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \a
f0 \loch \af0 {\*\bkmkstart article1.body1.sec4.sec2.p1}{\*\bkmkend article1.bod
y1.sec4.sec2.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe
2052 \langfenp2052 Blood collected from healthy and HIV-infected volunteers was
subjected to density gradient centrifugation using histopaque (Sigma) to separat
e plasma, red blood cells (RBC), and peripheral blood mononuclear cells (PBMC).
PBMCs were further processed to isolate monocytes and T cells. Plasma collected
was used for measurement of cytokines (IL-12, IL-2, IFN-\uc1 \u947 ?, IL-10 and
TNF-\uc1 \u945 ?) and free radicals.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loc
h \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \sb210
\lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21
\kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Isolation of human monocytes}{
\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfen
p2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \
rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart artic
le1.body1.sec4.sec3.p1}{\*\bkmkend article1.body1.sec4.sec3.p1}{\fs21 \kerning2
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 PBMCs isolated
from the blood of healthy volunteers and individuals with HIV infection were di
stributed into Poly-L-Lysine (Sigma) coated 96-well plates (1\uc1 \u215 ?10}{\fs
super1 5}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \l
angfenp2052 /well) and incubated overnight at 37\uc1 \u176 ?C, 5% CO}{\fs21 \ker
ning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \sub2 2}
52 , to allow monocytes to adhere. Non-adherent cells were removed and used for
isolation of T cells.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \
langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \sb210 \lisb0 \sa210 \
lisa0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0
\cgrid1 \langfe2052 \langfenp2052 Purification of T lymphocytes}{\b1 \fs24 \ker
ard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \f
s21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec4
.sec4.p1}{\*\bkmkend article1.body1.sec4.sec4.p1}{\fs21 \kerning2 \dbch \af2 \hi

ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Autologous T lymphocytes wer
e isolated from non-adherent cells (from both healthy volunteers and individuals
with HIV infection) using nylon wool columns (Polysciences). T cells were subje
cted to various treatments such as treatment with a GSH enhancing agent, N-acety
l cysteine (NAC) and treatment with a GSH synthesis inhibitor, buthionine sulpho
ximine (BSO). Specifically, T cells were treated as follows: no additives, treat
ment with NAC (5 mM), NAC (10 mM), NAC (20 mM), and treatment with BSO (500 \uc1
\u181 ?M), for 24 h. T cells were then washed, resuspended in fresh media conta
ining no additives and used for various assays such as measurement of GSH, deter
mination of the intracellular viability of }{\i1 \fs21 \kerning2 \dbch \af2 \hic
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 inside monocytes an
d cytokine production.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1
\langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \sb210 \lisb0 \sa210
\lisa0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbc
h \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f
0 \cgrid1 \langfe2052 \langfenp2052 Assay of GSH levels in T cells from healthy
and HIV-infected subjects}{\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \
ch \af0 {\*\bkmkstart article1.body1.sec4.sec5.p1}{\*\bkmkend article1.body1.sec
4.sec5.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \
langfenp2052 GSH levels were measured in freshly isolated T cells from healthy s
ubjects and individuals with HIV infection. Intracellular levels of GSH in T cel
ls were determined by spectrophotometry using an assay kit from Calbiochem. Brie
fly, T cells (2\uc1 \u215 ?10}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \
cgrid1 \langfe2052 \langfenp2052 \super1 5}{\fs21 \kerning2 \dbch \af2 \hich \af
0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 /well) were pelleted by centrifuga
tion and an equal volume of ice cold 5% metaphosphoric acid (MPA) was added to t
he pellet. Supernatants collected after centrifugation were analyzed for total G
SH as per manufacturer\uc1 \u8217 ?s instructions. Total GSH in the samples was
normalized with protein. Proteins in the samples were estimated by Bradford\uc1
\u8217 ?s method using Thermo Scientific Coomassie Protein Assay Reagent.}{\fs21
r }\pard \plain \qj \li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \
adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0
{\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfe
np2052 Assay of GSH levels in NAC/BSO-treated T cells from healthy and HIV-infec
ted subjects}{\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langf
e2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjus
tright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\b
kmkstart article1.body1.sec4.sec6.p1}{\*\bkmkend article1.body1.sec4.sec6.p1}{\f
We tested the effects of NAC and BSO in increasing and decreasing the intracellu
lar levels of GSH respectively, in T cells isolated from both healthy subjects a
nd individuals with HIV infection. T cells (2\uc1 \u215 ?10}{\fs21 \kerning2 \db
ch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \super1 5}{\fs21
\kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 /wel
l) isolated from healthy and HIV positive subjects were treated as follows: mock
treatment, treatment with NAC (10 mM) and treatment with BSO (500 \uc1 \u181 ?M
). Following overnight incubation, T cells were pelleted and used for GSH measur
ement as per manufacturer\uc1 \u8217 ?s instructions.}{\fs21 \kerning2 \dbch \af
\li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \lin0 \r
in0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Assay of free
radicals and TNF-\uc1 \u945 ? in plasma samples}{\b1 \fs24 \kerning2 \dbch \af2
\hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \
li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \d
bch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec4.sec7.p1}{\*\bkm

kend article1.body1.sec4.sec7.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f
0 \cgrid1 \langfe2052 \langfenp2052 Free radical levels in plasma samples derive
d from healthy subjects and individuals with HIV infection was determined by mea
suring the levels of malondialdehyde (MDA) using a colorimetric assay kit from C
ayman. Levels of TNF-\uc1 \u945 ? in plasma samples derived from healthy subject
s and individuals with HIV infection were determined by enzyme linked immuno-sor
bent assay (ELISA) using assay kits from eBioscience.}{\fs21 \kerning2 \dbch \af
\li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \lin0 \r
in0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2
\dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Preparation o
f bacterial cells for monocyte infection}{\b1 \fs24 \kerning2 \dbch \af2 \hich \
\hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec4.sec8.p1}{\*\bkmkend art
icle1.body1.sec4.sec8.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid
1 \langfe2052 \langfenp2052 All infection studies were performed using the virul
ent laboratory strain of }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \
cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \l
och \f0 \cgrid1 \langfe2052 \langfenp2052 , H37Rv inside the biosafety level 3 (
BSL-3) facility. }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \
langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0
\cgrid1 \langfe2052 \langfenp2052 was processed for infection as follows: stati
c cultures of H37Rv at their peak logarithmic phase of growth (between 0.5 and 0
.8 at A600) were used for infection of monocytes. The bacterial suspension was w
ashed and re-suspended in RPMI (Sigma) containing AB serum (Sigma). Bacterial cl
umps were disaggregated by vortexing five times with 3 mm sterile glass beads. T
he bacterial suspension was passed through a 5 \uc1 \u181 ?m syringe filter (Mil
lipore) to remove any further clumps. The total number of organisms in the suspe
nsion was ascertained by plating. Processed H37Rv was frozen as stocks at -80\uc
1 \u176 ?C. At the time of infection, frozen stocks of processed H37Rv were thaw
ed and used for monocyte infection.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \sb210 \
lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \
kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Co-incubation of }{\b1 \i1 \fs2
tb}{\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \la
ngfenp2052 -infected monocytes with autologous T cells}{\b1 \fs24 \kerning2 \dbc
\qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerni
ng2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec4.sec9.p1}{
\*\bkmkend article1.body1.sec4.sec9.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \l
och \f0 \cgrid1 \langfe2052 \langfenp2052 Adherent monocytes were infected with
processed H37Rv at a multiplicity of infection of 10\uc1 \u8758 ?1 and incubated
for 2 hours for phagocytosis. Unphagocytosed mycobacteria were removed by washi
ng the infected monocytes three times with sterile PBS. Infected monocytes were
cultured in RPMI containing 5% AB serum in presence and absence of autologous T
cells. Prior to co-incubation with infected-monocytes, autologous T cells were i
ncubated overnight with stimulants (NAC/BSO), washed with PBS, re-suspended in f
resh RPMI containing AB serum (without any stimulants) and then added to the inf
ected monocytes (monocyte: T cell ratio was adjusted to 1\uc1 \u8758 ?1). Infect
ed monocyte-T cell co-cultures were terminated at 1 hour and 5 days post-infecti
on to determine the intracellular survival of H37Rv. Infected monocytes cultured
in the absence of T cells were used as negative controls.}{\fs21 \kerning2 \dbc
\qj \li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \li
n0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \ker
ning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Terminat
ion of infected monocytes-autologous T cell co-cultures}{\b1 \fs24 \kerning2 \db

n \qj \li0 \ri0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap0 \fs21 \kern
ing2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec4.sec10.p1
}{\*\bkmkend article1.body1.sec4.sec10.p1}{\i1 \fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \a
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 -infected monocytes cu
ltured in the presence and absence of T lymphocytes were terminated at 1 hour an
d 5 days post-infection. During termination, supernatants were removed and adher
ent monocytes were lysed by addition of 200 \uc1 \u181 ?l sterile distilled wate
r. 25 \uc1 \u181 ?l of 10-fold diluted lysates were plated on 7H11 medium (Hi Me
dia) enriched with albumin dextrose complex (ADC), to estimate the extent of H37
Rv growth or killing in co-cultures of monocytes and T cells. Cell-free supernat
ants were plated to determine extracellular H37Rv growth. Cell-free supernatants
were also used for determining the levels of IL-12, IL-2 and IFN-\uc1 \u947 ?.}
52 \par }\pard \plain \qj \li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspa
lpha \adjustright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch
\af0 {\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \
langfenp2052 Assay of cytokines in supernatants derived from T cell-monocyte cocultures}{\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe205
2 \langfenp2052 \par }\pard \plain \qj \li0 \ri0 \widctlpar \aspalpha \adjustrig
ht \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmks
tart article1.body1.sec4.sec11.p1}{\*\bkmkend article1.body1.sec4.sec11.p1}{\fs2
1 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Ce
ll free-supernatants from co-cultures of }{\i1 \fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 M. tb}{\fs21 \kerning2 \dbch \a
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 -infected monocytes an
d T cells (from healthy controls and individuals with HIV infection) collected a
t 5 days post-infection were filtered using 0.2 micron syringe filters (Millipor
e) and assayed for the levels of IL-12, IL-2 and IFN-\uc1 \u947 ?. Cytokines wer
e assayed using ELISA kits from eBioscience.}{\fs21 \kerning2 \dbch \af2 \hich \
\sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustright \lin0 \rin0 \itap
0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \fs24 \kerning2 \dbch \a
f2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Assay of IL-12, IL-2,
IFN-\uc1 \u947 ? and IL-10 in plasma samples}{\b1 \fs24 \kerning2 \dbch \af2 \hi
\af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.body1.sec4.sec12.p1}{\*\bkmken
d article1.body1.sec4.sec12.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0
\cgrid1 \langfe2052 \langfenp2052 Levels of IL-12, IL-2, IFN-\uc1 \u947 ? and IL
-10 in plasma samples derived from healthy subjects and individuals with HIV inf
ection were determined by ELISA using assay kits from eBioscience.}{\fs21 \kerni
d \plain \qj \li0 \ri0 \sb210 \lisb0 \sa210 \lisa0 \widctlpar \aspalpha \adjustr
ight \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\b1 \f
Statistical Analysis}{\b1 \fs24 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid
f0 {\*\bkmkstart article1.body1.sec4.sec13.p1}{\*\bkmkend article1.body1.sec4.se
c13.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lan
gfenp2052 Statistical analysis of the data was carried out using Prism and Statv
iew\uc1 \u174 ? and the statistical significance was determined using an unpaire
d }{\i1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \lan
gfenp2052 t}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052
\langfenp2052 -test. Differences were considered significant at a level of P <0
.05.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langf
enp2052 \par }\pard \plain \qj \li0 \ri0 \sb174 \lisb0 \sa174 \lisa0 \widctlpar
\loch \af0 {\*\bkmkstart ack}{\*\bkmkend ack}{\b1 \fs31 \kerning2 \dbch \af2 \hi
ch \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 Acknowledgments}{\b1 \fs31 \
\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart article1.back1.a
ck1.p1}{\*\bkmkend article1.back1.ack1.p1}{\fs21 \kerning2 \dbch \af2 \hich \af0
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 We thank the staff at the Western U
niversity medical center for their help with blood draw. We thank the volunteers
for their participation in this study.}{\fs21 \kerning2 \dbch \af2 \hich \af0 \
21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\bkmkstart authcontrib}{\*\bkmk
end authcontrib}{\b1 \fs31 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \la
ngfe2052 \langfenp2052 Author Contributions}{\b1 \fs31 \kerning2 \dbch \af2 \hic
d1 \langfe2052 \langfenp2052 Conceived and designed the experiments: VV. Perform
ed the experiments: VV CG DM AS SK DG MT MF. Analyzed the data: VV CG DM. Contri
buted reagents/materials/analysis tools: FTK FG. Wrote the paper: VV.}{\fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\
pard \plain \qj \li0 \ri0 \sb174 \lisb0 \sa174 \lisa0 \widctlpar \aspalpha \adju
stright \lin0 \rin0 \itap0 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\*\
bkmkstart references}{\*\bkmkend references}{\b1 \fs31 \kerning2 \dbch \af2 \hic
h \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 References}{\b1 \fs31 \kernin
\plain \qj \fi0 \li360 \ri0 \widctlpar \aspalpha \adjustright \lin360 \rin0 \tq
l \tx720 \itap0 \ls2 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 1. {\*\
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ngfe2052 \langfenp2052 \cf13 \ul1 View Article}}}{\fs21 \kerning2 \dbch \af2 \hi
2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard
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ql \tx1440 \tql \tldot \tx2357 \itap0 \ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hi
ch \af0 \loch \af0 {\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hic
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ponses+in+persons+with+tuberculosis+and+human+immunodeficiency+virus+infection."
}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2
cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 PubMed/NCBI}}}{\fs21 \kerning2 \dbch
r }\pard \plain \qj \fi0 \li1080 \ri0 \widctlpar \aspalpha \adjustright \lin1080
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+responses+in+persons+with+tuberculosis+and+human+immunodeficiency+virus+infecti
on.&author=Zhang&publication_year=1994"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2
l1 Google Scholar}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \l
widctlpar \aspalpha \adjustright \lin360 \rin0 \tql \tx720 \itap0 \ls2 \fs21 \ke
rning2 \dbch \af2 \hich \af0 \loch \af0 {\fs21 \kerning2 \dbch \af2 \hich \af0 \
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\*\bkmkend pone.0028378-Buhl1}Buhl R, Jaffe HA, Holroyd KJ, Wells FB, Mastrangel
i A, et al. (1989) Systemic glutathione deficiency in symptom-free HIV seroposit
ive individuals. Lancet 2: 1294\uc1 \u8211 ?8. }{\fs21 \kerning2 \dbch \af2 \hic
li1080 \ri0 \widctlpar \aspalpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \
tldot \tx2357 \itap0 \ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
0 {\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://d
x.doi.org/10.1016/s0140-6736(89)91909-0"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af
ul1 View Article}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \la
widctlpar \aspalpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \tldot \tx2357
\itap0 \ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\field \fl
f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://www.ncbi.nlm.n
ih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Tit
le+Word&term=Buhl[author]+AND+Systemic+glutathione+deficiency+in+symptom-free+HI
V+seropositive+individuals."}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0
\af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 PubMed/N
CBI}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \lan
langfe2052 \langfenp2052 \par }\pard \plain \qj \fi0 \li1080 \ri0 \widctlpar \as
palpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \tldot \tx2357 \itap0 \ls1
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lookup?title=Systemic+glutathione+deficiency+in+symptom-free+HIV+seropositive+in
dividuals.&author=Buhl&publication_year=1989"}}{\*\fldrslt {\fs21 \kerning2 \dbc
f13 \ul1 Google Scholar}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgr
\loch \f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \fi0 \li360
\ri0 \widctlpar \aspalpha \adjustright \lin360 \rin0 \tql \tx720 \itap0 \ls2 \fs
21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fs21 \kerning2 \dbch \af2 \hich
\af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 4. {\*\bkmkstart pone.0028378-S
taal1}{\*\bkmkend pone.0028378-Staal1}Staal F, Roederer M, Herzenberg LA, Herzen
berg LA (1990) Intracellular thiols regulate activation of nuclear factor \uc1 \
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id1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \fi0 \li1080 \ri0 \widctlpa
r \aspalpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \tldot \tx2357 \itap0
\ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\field \fldedit {\
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s.87.24.9943"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cg
0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 View Article}}}{\fs21 \
ngfenp2052 \par }\pard \plain \qj \fi0 \li1080 \ri0 \widctlpar \aspalpha \adjust
right \lin1080 \rin0 \tql \tx1440 \tql \tldot \tx2357 \itap0 \ls1 \ilvl1 \fs21 \
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AND+Intracellular+thiols+regulate+activation+of+nuclear+factor+?B+and+transcript
ion+of+human+immunodeficiency+virus."}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \
PubMed/NCBI}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe
\cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \fi0 \li1080 \ri0 \widc
tlpar \aspalpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \tldot \tx2357 \it
ap0 \ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\field \fldedi
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/scholar_lookup?title=Intracellular+thiols+regulate+activation+of+nuclear+factor
+?B+and+transcription+of+human+immunodeficiency+virus.&author=Staal&publication_
year=1990"}}{\*\fldrslt {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid
loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 Google Scholar}}}{\fs21 \k
gfenp2052 \par }\pard \plain \qj \fi0 \li360 \ri0 \widctlpar \aspalpha \adjustri
ght \lin360 \rin0 \tql \tx720 \itap0 \ls2 \fs21 \kerning2 \dbch \af2 \hich \af0
\loch \af0 {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052
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l2}Staal FJ (1998) Glutathione and HIV infection: reduced reduced, or increased
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h \af2 \hich \af0 \loch \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 PubMed/
NCBI}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \langfe2052 \la
\langfe2052 \langfenp2052 \par }\pard \plain \qj \fi0 \li1080 \ri0 \widctlpar \a
spalpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \tldot \tx2357 \itap0 \ls1
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ri0 \widctlpar \aspalpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \tldot \t
x2357 \itap0 \ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\fiel
och \f0 \cgrid1 \langfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://dx.doi.or
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i?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Hirsch[aut
hor]+AND+Depressed+T+cell+interferon-gamma+responses+in+pulmonary+tuberculosis:+
analysis+of+underlying+mechanisms+and+modulation+with+therapy."}}{\*\fldrslt {\f
2 \langfenp2052 \cf13 \ul1 PubMed/NCBI}}}{\fs21 \kerning2 \dbch \af2 \hich \af0
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np2052 \cf13 \ul1 View Article}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \a
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\tldot \tx2357 \itap0 \ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \a
f0 {\field \fldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af
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\ul1 PubMed/NCBI}}}{\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \l
\f0 \cgrid1 \langfe2052 \langfenp2052 \par }\pard \plain \qj \fi0 \li1080 \ri0
\widctlpar \aspalpha \adjustright \lin1080 \rin0 \tql \tx1440 \tql \tldot \tx235
7 \itap0 \ls1 \ilvl1 \fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 {\field \f
ldedit {\*\fldinst {\fs21 \kerning2 \dbch \af2 \hich \af0 \loch \af0 \cgrid1 \la
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ari1}Leghmari K, Bennasser Y, Tkaczuk J, Bahraoui E (2008) HIV-1 Tat protein ind
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ngfe2052 \langfenp2052 \cf13 \ul1 HYPERLINK "http://dx.doi.org/10.1016/j.cellimm
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t pone.0028378-Jaruga1}{\*\bkmkend pone.0028378-Jaruga1}Jaruga P, Jaruga B, Gack
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ive modification of DNA in lymphocytes of HIV-infected patients. Free Radic Biol
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bmeyer1}{\*\bkmkend pone.0028378-Dobmeyer1}Dobmeyer TS, Findhammer S, Dobmeyer J
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Gluthatione and Immunity

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sponses, as evidenced by decreased production of the cytokines IL-2 and IFN-\uc1

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HIV infected individuals compared to healthy subjects (}{\field \fldedit {\*\fld

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52 \langfenp2052 infection. We also quantified the levels of IL-12, IL-2 and IF

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son ME, Sharma VK, Meister A (1990) Glutathione regulates activation-dependent D

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