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En el caso de la India, el budismo se ha ramificado en multitud de escuelas.

In this chapter I established the bead-based assays to measure the phagocytosis ability

of macrophage as well as its antimicrobial activities including acidification and

proteolysis. At first I successfully generated the beads coated with ligands and

line to use it as a model to set up and optimize the bead-based assays. In such models,

the assays were shown to work well giving consistent results. I observed the decrease in

phagosomal pH and the occurrence of proteolysis in macrophage treated with beads. I

found that stimulation of macrophages with TDM or β-glucan led to a significantly

decreases in proteolytic activities in comparison to resting macrophage. Interaction

between macrophages and bacteria could be investigated by infection of macrophages

with Mtb expressing mCherry. Our assays also worked in the hMDM model which

displayed variations in antimicrobial activities of macrophages from different subjects.

The bead-based assays were originally developed to study the physiological changes in

phagosome of macrophage that can help to explain its plastic functions in both

homeostasis and during immune response. Therefore, it is suggested that this new

approach can be applied in clinical settings to identify the macrophage-associated

pathologies275,273,226,299. The bead-based measurement of antimicrobial activities of AM

in HIV patients have suggested the possible association of the impairment in AM

activities and the increasing susceptibility to upper respiratory disease274. Furthermore,

this new approach has helped to provide the insight to the global modification in

hMDM activities in response to Mtb infection239.

The beads used in these studies were only opsonized with IgG to facilitate the uptake by

macrophages. It remains unclear whether different routes of entry into macrophages

may lead to differences in subsequent events within the host such as signal transduction,

immune activation, and intracellular survival of Mtb as well as outcome of infection14.

The beads developed in our study were novel since they were coated with specific

contribution of the signaling pathways specific for these ligands in controlling

macrophage antimicrobial activities in response to Mtb infection.

By exploiting the beads labeled with pH-sensitive fluorochrome carboxyfluorescein SE,

I have seen a rapid redution in phagosomal pH, which had been observed in previous

studies239, from neutral to 5.5 within 1 h in resting macrophage. The bulk protease

activity of macrophages was assessed by the beads coupled with DQ green BSA. I

observed an increase in fluorescence signal generated by degradation of this substrate

overtime, indicating the occurrence of proteolysis in phagosome. J774 macrophages

treated with TDM- or β-glucan- coated beads showed a reduced proteolysis activity

compared with resting macrophage. The beads are coated with an excess amount of

ligands (IgG/TDM or β-glucan) so that they are optimal for recognition and phagocytosis which is further indicated by the comparable phagocytic ability of

macrophages towards IgG, TDM or β-glucan beads. Therefore, the variations in proteolysis activity of macrophages treated with different ligands-coated beads could

come from the nature of ligands. Different ligands mediate their specific pathways that

would induce different macrophage responses. The decreased activity in macrophages

treated with TDM- or β-glucan beads was similar to that observed previously upon

activation of macrophages with LPS272. These results suggest a shift in macrophage

function from its role in homeostasis to the role in immunity. More clearly, the

homeostatic function of macrophages is to remove dead cells or cell debris, hence their

hydrolytic activity is programmed to occur with high efficiency for complete

degradation of those materials. Whereas, under activation by cytokines or bacterial

and efficient antigen processing for presenting to T cells. The consistency in our data

and previous data also provided a validation for the reliability of our bead-based assays.

The Mtb strain expressing mCherry provided a sensitive and rapid tool in a study of

macrophage-mycobacterial interaction. As other reporter Mtb strains generated in

previous study, our strain showed a strong correlation between mCherry intensity and

bacteria viability276. mCherry signal was very sensitive, so the signal could be detected

at a very low cell density of Mtb. It has been revealed that the fate of Mtb-infected

macrophage is different due to the intracellular burden of bacteria300, cells with heavily

burden progressing to necrosis. The reporter fluorescence in our study facilitated the

assessment the correlation of intracellular growth of Mtb and the macrophage lysis

when bacteria were infected to murine and human macrophage cell lines at different

MOI. In consistent with the previous study290, I also observed the rapid increase of

intracellular Mtb growth at MOI 2 and 3 that leads to early host cell lysis in comparison

to MOI 1 in both J774 and THP1 models. Because of slow bacterial growth and delay in

host cell lysis, I propose that the infection of macrophage at MOI 1 is applicable for

examining the host-pathogen interactions in detail.

In summary I have successfully established the assays to examine the phagosomal

antimicrobial activities as well as killing ability of macrophages; these assays showed

consistent results during experimental replications. In the next chapter, I apply these

assays developed here to study the association of macrophage antimicrobial activities