Abstract
Several microorganisms are currently being used as production platform for glycolipid biosurfactants, providing a greener alternative to chemical biosurfactants. One of the reasons why these processes are commercially competitive is the fact that microbial producers can efficiently export their product to the extracellular environment, reaching high product titers. Glycolipid biosynthetic genes are often found in a dedicated cluster, amidst which genes encoding a dedicated transporter committed to shuttle the glycolipid to the extracellular environment are often found, as is the case for many other secondary metabolites. Knowing this, one can rely on gene clustering features to screen for novel putative transporters, as described and performed in this review. The above strategy proves to be very powerful to identify glycolipid transporters in fungi but is less valid for bacterial systems. Indeed, the genetics of these export systems are currently largely unknown, but some hints are given. Apart from the direct export of the glycolipid, several other transport systems have an indirect effect on glycolipid production. Specific importers dictate which hydrophilic and hydrophobic substrates can be used for production and influence the final yields. In eukaryotes, cellular compartmentalization allows the assembly of glycolipid building blocks in a highly specialized and efficient way. Yet, this requires controlled transport across intracellular membranes. Next to the direct export of glycolipids, the current state of the art regarding this indirect involvement of transporter systems in microbial glycolipid synthesis is summarized in this review.
Key points
• Transporters are directly and indirectly involved in microbial glycolipid synthesis.
• Yeast glycolipid transporters are found in their biosynthetic gene cluster.
• Hydrophilic and hydrophobic substrate uptake influence microbial glycolipid synthesis.
Graphical abstract

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Acknowledgements
The authors wish to thank the Research Foundation - Flanders (FWO; grant number 28930 and 1185417N), as well as the Special Research Fund of Ghent University Ghent (BOF; grant number STA011-17) for financial support.
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SC screened the literature and wrote the manuscript. LJS collaborated in overall writing and wrote on glycerol transport mechanisms. INAVB was involved in writing part of the text on CBL, conception, guiding, and reviewing. All authors read and approved the manuscript.
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Claus, S., Jenkins Sánchez, L. & Van Bogaert, I.N.A. The role of transport proteins in the production of microbial glycolipid biosurfactants. Appl Microbiol Biotechnol 105, 1779–1793 (2021). https://doi.org/10.1007/s00253-021-11156-7
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DOI: https://doi.org/10.1007/s00253-021-11156-7
Keywords
- Biosurfactant
- Glycolipid
- Transporter
- Gene cluster
- Yeast
- Bacteria