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antibodies generated against proteins <strong>of</strong> <strong>the</strong> endomembrane system showed that cyst<br />

wall proteins were exported through <strong>the</strong> re-organisation membrane compartments<br />

that were biochemically similar to ESVs (Marti et al. 2003). These results<br />

demonstrate ESVs are part <strong>of</strong> a Golgi cisternae-like system which is constitutively<br />

expressed ra<strong>the</strong>r than being generated de novo.<br />

Excystation<br />

Within <strong>the</strong> mammalian host, excystation occurs rapidly after passage through <strong>the</strong><br />

acidic environment <strong>of</strong> <strong>the</strong> stomach <strong>and</strong> after entry into <strong>and</strong> exposure to elements <strong>of</strong><br />

<strong>the</strong> proximal small intestine (Adam 2001). Excystation was first induced in vitro<br />

using acidic solutions (optimum pH 1.3 – 2.7; Bingham & Meyer 1979) which<br />

suggest an importance <strong>of</strong> passage through <strong>the</strong> acidic environment <strong>of</strong> <strong>the</strong> stomach for<br />

excystation. This may not always be <strong>the</strong> case, with G. muris excystation having<br />

occurred in vitro at pH 7.5 in vitro (Feely, Gardner <strong>and</strong> Hardin 1991) <strong>and</strong> may<br />

explain cases <strong>of</strong> giardiasis in patients with reduced gastric acidity. Subsequently a pH<br />

<strong>of</strong> 4 has been assessed as <strong>the</strong> optimum pH <strong>and</strong> that <strong>the</strong> use <strong>of</strong> pancreatic proteases<br />

such as trypsin or chymotrypsin being required for in vitro excystation <strong>and</strong> suggest<br />

<strong>the</strong> role for pancreatic secretions for excystation in vivo (Boucher & Gillin 1990).<br />

Life cycle<br />

Humans are indicated as reservoirs <strong>of</strong> this protozoan, although o<strong>the</strong>r mammals<br />

including dogs, cats, farm animals <strong>and</strong> beavers have been shown to harbour Giardia<br />

(Ortega & Adam 1997; McGlade et al. 2003; Hunter & Thompson 2005).<br />

Transmission in humans may occur via direct faecal-oral contact or from faecally<br />

contaminated water <strong>and</strong> food (Smith et al. 2007). The zoonotic transmission <strong>of</strong><br />

Giardia is still unclear but has been described in nor<strong>the</strong>ast India from dogs (Traub et<br />

al. 2004) <strong>and</strong> direct contact with farm animals <strong>and</strong> pets have also been associated<br />

with giardia infection (Warburton, Jones <strong>and</strong> Bruce 1994; Hunter & Thompson<br />

2005).<br />

The infection cycle begins with <strong>the</strong> consumption <strong>of</strong> infective cysts, usually from <strong>the</strong><br />

faecal-oral route by contaminated food or water (Figure 1.5). After <strong>the</strong> safe passage<br />

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