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LEISHMANIA 231<br />

(1,6)myo-inositol-1-PO 4 -lipid motif, the hallmark<br />

of all GPI anchors. LPPG differs from<br />

the Leishmania GIPLs by the presence of the<br />

phosphoceramide moiety, a feature found<br />

in the LPPG of Acanthamoeba castellanii and<br />

in a lipid anchor found in Dictyostelium<br />

discoideum. Moreover T. cruzi LPPG contains<br />

2-aminoethylphosphonate as does the<br />

Acanthamoeba castellanii phosphoglycan.<br />

LEISHMANIA<br />

Leishmania exist as flagellated, extracellular<br />

promastigotes in the midgut of their sand fly<br />

vector, and as intracellular, aflagellar amastigotes<br />

in the phagolysosomes of the mammalian<br />

macrophage. The parasites are <strong>trans</strong>mitted<br />

when a sand fly takes a blood meal from an<br />

infected individual. Within the sand fly midgut<br />

the promastigotes develop from the avirulent<br />

procyclic form to the virulent metacyclic<br />

form in a process termed metacyclogenesis.<br />

Throughout their life cycle, Leishmania survive<br />

and proliferate in highly hostile environments<br />

and have evolved special mechanisms<br />

which enable them to endure these adverse<br />

conditions, including a dense cell surface glycocalyx<br />

composed of lipophosphoglycan,<br />

glycosylinositolphospholipids and secreted<br />

glycoconjugates, proteophosphoglycan (PPG)<br />

and secreted acid phosphatase (sAP).<br />

Lipophosphoglycan<br />

Lipophosphoglycan (LPG), the predominant<br />

cell surface glycoconjugate of Leishmania<br />

promastigotes, is localized over the entire parasite<br />

surface including the flagellum. Found in<br />

all species of Leishmania that infect humans,<br />

sAP<br />

LPG<br />

PG<br />

Cap<br />

Repeating unit<br />

GIPLs<br />

GPI-anchored<br />

proteins<br />

PPG<br />

-Ser-PO 4 -<br />

Core<br />

FIGURE 10.3 Schematic representation of the surface coat and secreted glycoconjugates of Leishmania. The small<br />

oval ‘repeat unit’ represents Gal-Man-PO 4 units. In PPG and sAP, the repeat unit chain is attached to the polypeptide<br />

via Man-PO 4 -serine linkage. GIPLs, glycosylinositolphospholipids; LPG, lipophosphoglycan; PG, phosphoglycan;<br />

PPG, proteophosphoglycan; sAP, secreted acid phosphatase.<br />

BIOCHEMISTRY AND CELL BIOLOGY: PROTOZOA

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