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150 ENERGY METABOLISM – TRYPANOSOMATIDAE<br />

Oxidative<br />

pathway<br />

Glucose 6-phosphate<br />

NADP H 2 O<br />

1<br />

NADPH 2H <br />

6-Phosphogluconolactone<br />

2 H 2 O<br />

6-Phosphogluconate<br />

NADP <br />

3<br />

NADPH H <br />

CO 2<br />

Ribulose 5-phosphate<br />

4 5<br />

Xylulose<br />

5-phosphate<br />

6<br />

Ribose<br />

5-phosphate<br />

Non-oxidative<br />

pathway<br />

8<br />

Sedoheptulose<br />

7-phosphate<br />

Erythrose<br />

4-phosphate<br />

7<br />

Glyceraldehyde<br />

3-phosphate<br />

Fructose<br />

6-phosphate<br />

Fructose<br />

6-phosphate<br />

Glyceraldehyde<br />

3-phosphate<br />

FIGURE 7.5 The hexose-monophosphate pathway. Enzymes: (1) Glucose-6-phosphate dehydrogenase;<br />

(2) 6-phosphogluconate lactonase; (3) 6-phosphogluconate dehydrogenase; (4) ribulose 5-phosphate 3-epimerase;<br />

(5) ribulose 5-phosphate isomerase; (6) <strong>trans</strong>ketolase; (7) <strong>trans</strong>aldolase; (8) <strong>trans</strong>ketolase.<br />

pathway may be associated with glycosomes<br />

as well. While 50% of the activity of glucose-6-<br />

phosphate dehydrogenase (G6PDH) in T. brucei<br />

is found in glycosomes, its inferred protein<br />

sequence did not reveal a peroxisome-targeting<br />

signal. However, the T. brucei 6-phosphogluconolactonase<br />

(6PGL), of which some 15% of<br />

the activity is glycosomal, contains a C-terminal<br />

motif showing similarity to the established<br />

type of targeting signal, in agreement with its<br />

subcellular localization. The corresponding<br />

genes from L. mexicana have also been cloned<br />

and sequenced and here the G6PDH contains<br />

a PTS 1 import signal, while the 6PGL did not.<br />

However, the sequence of the third enzyme in<br />

the pathway, 6-phosphogluconate dehydrogenase<br />

(6PGDH), does not contain a detectable<br />

PTS. Interestingly, the sequences of 6PGL and<br />

6PGDH in both organisms are of prokaryotic<br />

origin and therefore most likely entered the<br />

BIOCHEMISTRY AND CELL BIOLOGY: PROTOZOA

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