Rhodococci are able to produce TAG from a wide diversity
of carbon sources; including mono- and disac-Biosynthesis of TAG requires an efficient metabolic
network able to produce the necessary precursors, reducing
power and energy for the specific reactions. In
general, the pathways of rhodococcal central metabolism
are able to efficiently convert diverse carbon
sources to key metabolic intermediates, such as pyruvate,
acetyl-CoA and glycerol-3-phosphate, to create
reducing equivalents that are required by lipid biosynthesis
pathways and to produce the necessary energy
as ATP. The conversion of acetyl-CoA from glycolysis
-derived pyruvate might be the major route of carbon
flux to fatty acid biosynthesis by some rhodococci;
thus, sugars and related substrates support significant
TAG accumulation by Rhodococcus members (Figure
1A). However, the total TAG content may vary noticeably
among individual species or strains within
rhodococci, and are ultimately controlled by the genetic
make-up of individual organisms (Figure 1A and
B). The capacity of rhodococcal cells for the uptake
and catabolism/assimilation of the available carbon
source seems to also be relevant for TAG accumulation.
For instance, the biosynthesis of TAG from gluconate/
glucose and hexadecane involves different
metabolic pathways (Figure 3). When cells are cultivated
on gluconate or glucose (unrelated carbon
sources), the substrate has to be degraded to acetyl-
CoA, which serves as precursor for fatty acid biosynthesis.
In contrast, during cultivation with hexadecane
(related carbon source) the cells utilize the preformed
fatty acids derived from the mono-terminal oxidation
of the alkane for TAG biosynthesis (Figure 3) (Alvarezet al. 1997, Alvarez 2003).
Depending on the substrate used as the carbon source, more than one pathway of central metabolism may contribute to producing and maintaining the ace-tyl-CoA pool available for fatty acid and TAG biosyn-thesis. The production of the key metabolic precursors for TAG biosynthesis, the fatty acids, is performed in rhodococci by a multienzymatic complex known as fatty acid synthase I (FASI). This complex catalyzes the successive reaction of condensation, reduction, dehydration and reduction. The FAS type I consists in a unique large protein with different catalytic activi-ties. FASI enzymes are found in the cytoplasm of eu-karyotic cells and in a subgroup of actinobacteria, such as mycobacteria, which are used for phospholipids and TAG synthesis or for mycolic acid production after an elongation process mediated by FASII (Bloch 1977, Zimhony et al. 2004). The FASI multienzyme gene of mycobacteria and rhodococci seem to be structurally very similar. The main products of rhodococcal FASI may be C16- C18 fatty acids, which may be utilized for phospholipids and TAG biosynthesis. Fatty acids, probably as acyl-CoA residues, are sequentially incor-porated into the glycerol-3-phosphate backbone for TAG biosynthesis.