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184 PROTEIN METABOLISM<br />

PROLINE<br />

Polyamines (1) (2)<br />

(16)<br />

Ornithine<br />

(3)<br />

Glutamate semialdehyde<br />

(4)<br />

GLUTAMATE<br />

(5)<br />

(12)<br />

α-ketoglutarate<br />

∇<br />

-pyrroline-5-carboxylate<br />

GLUTAMINE<br />

(6)<br />

NH 3<br />

Succinate<br />

(8)<br />

Succinic<br />

semialdehyde<br />

ARGININE<br />

(13)<br />

Citrulline NH 3<br />

(14)<br />

Carbamoyl phosphate<br />

(15)<br />

NH 3 CO 2<br />

TCA cycle<br />

(7)<br />

γ-aminobutyrate<br />

Urea<br />

(11)<br />

(9)<br />

γ-guanidinobutyrate<br />

O 2<br />

(10)<br />

NH 3<br />

CO 2 γ-guanidinobutyramide<br />

FIGURE 8.2 Catabolic pathways of proline, glutamine, glutamate and arginine. (1) proline oxidase; (2) -pyrroline-<br />

5-carboxylate reductase; (3) non-enzymatic reaction; (4) glutamate semialdehyde dehydrogenase; (5) glutamate<br />

dehydrogenase; (6) glutaminase; (7) -aminobutyrate amino<strong>trans</strong>ferase; (8) succinic semialdehyde dehydrogenase;<br />

(9) arginine decarboxyoxidase; (10) -guanidinobutyramide aminohydrolase; (11) -guanidinobutyrate amidinohydrolase;<br />

(12) arginase; (13) arginine deiminase; (14) catabolic ornithine <strong>trans</strong>carbamoylase; (15) carbamate<br />

kinase; (16) ornithine amino<strong>trans</strong>ferase.<br />

T. vaginalis and T. foetus, ultimately leads to<br />

ammonia and carbon dioxide formation. This<br />

pathway is unusual in eukaryotes; it is usually<br />

restricted to prokaryotes. In trichomonads,<br />

ornithine <strong>trans</strong>carbamoylase and carbamate<br />

kinase are predominantly cytosolic, whereas<br />

the first enzyme of the pathway, arginine deiminase,<br />

is particulate. The arginase and the<br />

-guanidinobutyramide pathways lead to the<br />

generation of ATP under aerobic conditions via<br />

the TCA cycle. On the other hand, the dihydrolase<br />

pathway is operative under anaerobic conditions,<br />

and results in the direct formation of<br />

ATP in the carbamate kinase reaction. Indeed,<br />

the dihydrolase pathway appears to be responsible<br />

for a significant proportion of the ATP<br />

generated by these protozoa, although glucose<br />

remains as the major fuel, at least in the case<br />

of T. vaginalis. The enzymes of the dihydrolase<br />

pathway are present in G. lamblia at considerably<br />

higher levels than in T. vaginalis.<br />

The arginase pathway is operative in many<br />

helminths; in some trematodes, glutamate<br />

semialdehyde enters the biosynthetic pathway<br />

of proline, which is then the final product<br />

of arginine catabolism.<br />

BIOCHEMISTRY AND CELL BIOLOGY: PROTOZOA

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