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Mousley, C. The Secsuperfamily and the regulatory interface between phospholipid metabolism and membrane trafficking. Litvak, V. Maintenance of the diacylglycerol level in the Golgi apparatus by the Nir2 protein is critical for Golgi secretory function.
Chernomordik, L. Lipids in biological membrane fusion. Shemesh, T. Prefission constriction of Golgi tubular carriers driven by local lipid metabolism: a theoretical model. Gennis, R. Takamori, S. Molecular anatomy of a trafficking organelle. A careful reconstruction of synaptic vesicles shows that cholesterol and phospholipids molar ratio 0. Dietrich, C. Partitioning of Thy-1, GM1, and cross-linked phospholipid analogs into lipid rafts reconstituted in supported model membrane monolayers.
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Lagerholm, B. Detecting microdomains in intact cell membranes. Meder, D. Phase coexistence and connectivity in the apical membrane of polarized epithelial cells. Molecular dynamics and interactions for creation of stimulation-induced stabilized rafts from small unstable steady-state rafts.
Bollinger, C. Ceramide-enriched membrane domains. Roux, A. Role of curvature and phase transition in lipid sorting and fission of membrane tubules. Chiantia, S. Sot, J. Anishkin, A.
Searching for the molecular arrangement of transmembrane ceramide channels. Relationship of lipid rafts to transient confinement zones detected by single particle tracking. Sengupta, P. Dibble, A. Biochemistry 33 , — Lewis, R. Biochemistry 46 , — Jacobson, K. Lipid rafts: at a crossroad between cell biology and physics.
On receiving a signal, proteins control phase behaviour by combining their shell with similar lipid shells of other proteins. Shogomori, H. Palmitoylation and intracellular domain interactions both contribute to raft targeting of linker for activation of T cells.
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Polyunsaturated acyl chains of membrane lipids can effectively drive the formation of membrane rafts because of especially poor packing with cholesterol. Kucerka, N. Closer look at structure of fully hydrated fluid phase DPPC bilayers. Liu, K. Cell 18 , — Download references. You can also search for this author in PubMed Google Scholar. Correspondence to Gerrit van Meer. Lipid MAPS. Gerrit van Meer's homepage. Dennis R.
Voelker's homepage. Gerald W. Feigenson's homepage. A family of storage lipids consisting of glycerol esterified to three fatty acids, forming the hydrophobic core of lipid droplets and blood lipoproteins together with steryl esters. A family of storage lipids consisting of sterol esterified to one fatty acid, forming the hydrophobic core of lipid droplets and blood lipoproteins together with triacylglycerol molecules.
Dolicholphosphate and dolicholpyrophosphate anchor sugar molecules to the ER membrane for transfer to proteins in the ER lumen.
An endosome containing internal vesicles that originate from inward budding. This direction of budding is away from the cytosol opposite to the regular budding of transport vesicles and a different molecular machinery has been found to be responsible. One of a family of membrane-embedded transporters that share a phosphorylated intermediate as part of their reaction cycle. A mechanism involving incompletely characterized components that allows transmembrane movement of lipids and relaxation of lipid asymmetry on cell stimulation.
The coat protein of caveolae. It is anchored by a hydrophobic loop and 1—3 palmitoylated cysteines. Caveolin interacts with cholesterol and oligomerizes. Reprints and Permissions. Membrane lipids: where they are and how they behave.
Nat Rev Mol Cell Biol 9, — The synthesis of tetrahymanol, another pentacyclic structure related to HOPs, was also shown to be Shc-dependent in Rhodopseudomonas palustris Welander et al. Shc is an ortholog of squalene-2,3-epoxide cyclase responsible for the formation of lanosterol from oxidosqualene. A Shc homolog from firmicutes has been shown to be responsible for the synthesis of sporulenes Bosak, Losick and Pearson ; Kontnik et al. Synthesis of HOPs in bacteria.
Starting from diploptene elongated HOPs can be synthesized. Often, genes involved in HOP synthesis and modification form operons and cluster within the bacterial genomes. HpnG is responsible for the release of adenine from adenosylhopane Fig. HpnG shows homology to purine nucleoside phosphorylases, but there is still a debate if the reaction product is phosphoribohopane or ribosylhopane. It has been suggested that the cyclic ribose opens up without the aid of an enzyme Bradley et al.
Formylhopane is probably the substrate for the aminotransferase reaction catalyzed by HpnO Fig. The conversion of ribosylhopane to BHT has been also suggested to happen non-enzymatically, but it is unclear how the required reduction should be achieved this way Schmerk et al.
HpnI is a glycosyltransferase and HpnK has been suggested to be a deacetylase. HpnJ is a radical SAM superfamily member required for the formation of BHT cyclitolether, but the mechanism of the ring contraction reaction is not known Fig. HOPs can also be C6- and Cunsaturated, but the enzymes responsible for these unsaturation reactions have not been identified Fig. C2 and C3 methylation reactions of HOPs have been described and the methyltransferases have been identified Fig.
In contrast, 3-methylhopanoids have been used as a biomarker proxy for aerobic methanotrophs in the past, but it is clear now that several proteobacteria and actinomycetes also have a gene encoding homologs of the methyltransferase HpnR required for 3-methylhopanoid formation. Mutants in different steps of HOP formation have been characterized. Rhodopseudomonas palustris mutants deficient in HOP formation presented a weakened outer membrane and were more sensitive to pH and temperature stress Welander et al.
Burkholderia cenocepacia defective in HOP formation displayed increased sensitivity to low pH, detergents, various antibiotics and did not produce flagella Schmerk, Bernards and Valvano In contrast, in S. The phospholipase C SMc from S.
DAG can then be used in S. Interestingly, the PE methyltransferase from S. Genes encoding for SMclike enzymes are absent in intracellular pathogens such as Brucella , Bartonella and Rickettsia. This absence of a phosphate starvation response is probably the consequence of an intracellular lifestyle which is characterized by a stable phosphate supply. Bacteria change their membrane lipid composition in response to changes in the environment.
This adaptation allows bacteria to survive unfavorable conditions. Even during the simple experiment of growing microorganisms in a flask, the environment the bacteria are sensing is changing constantly, which will be affecting the membrane lipid composition: nutrient levels fall, products of metabolism accumulate, the pH of the culture medium may rise or fall and oxygen levels are changing.
The probably best-known example of membrane adaptation happens when bacteria are growing at different temperatures. An increase in temperature causes an increase in fluidity and probably an increase in the occurrence of discontinuities in the membrane.
Bacteria growing at increased temperature usually contain more saturated fatty acids in their membranes Marr and Ingraham However, membrane modifications as a response to abiotic stress conditions are not restricted to the fatty acid moieties and do also happen on headgroup level. The membrane lipid modifications occurring can be divided in two types. This modification of pre-existing membrane lipids has the advantage that it allows a quick response to changes in environmental conditions.
In the following, we will discuss how membrane lipid modifications occur under different stress conditions. Growth under acidic conditions provokes a modification in membrane lipids. The second gene of the operon encodes the putative lipase AtvA. This operon structure is also present in P.
Thus, it can be hypothesized that AtvA from R. This modification of the anionic membrane lipid PG to the cationic LPG or zwitterionic APG membrane lipids has been shown to be important in resistence to cationic peptides and antibiotics. Another lipid modification described in R. It is thought that the presence of the additional hydroxyl group allows the formation of hydrogen bonds that increase the lateral interactions between lipid molecules and stabilize the leaflet structure Nikaido Consistent with this hypothesis, R.
The 2-hydroxylation of OL also seems to play a role in the response to increased growth temperature in R. During the response to high osmotic pressures, the anionic membrane lipid CL has been shown to play an important role in different bacteria. Increased osmotic pressure causes an increase in CL formation in E. ProP is denoted an osmosensory transporter because it is activated by increasing osmolarity and causes the accumulation of the compatible solute proline allowing bacterial growth at increased osmotic pressure.
The activation threshold of ProP correlates with the amount of CL present in the membrane and its presence controls the amount and activity of the proline transporter ProP Romantsov et al.
Many soil bacteria remodel their membranes in response to conditions of phosphate limitation. Plants and microbes mostly live in environments where available phosphate is a growth-limiting factor.
Apparently, some bacteria can replace their phospholipids with membrane lipids devoid of phosphorus. This remodeling has been studied in some detail in R. It was suggested that phospholipids are functionally replaced by phosphorus-free membrane lipids under conditions where phosphate becomes limiting thereby making the phosphate pool present in the membrane accessible for other cellular processes such as nucleic acid synthesis.
There is evidence supporting these hypotheses: Recently, Riekhof and collaborators constructed a S. They showed that essential processes of membrane biogenesis and organelle assembly were functional in this strain Riekhof et al.
When an A. Bacteria account for most of the diversity of life on our planet and they have been around for billions of years. During this time they have adapted to all types of environments on Earth. From this point of view, it is probably not surprising that bacteria present such a rich diversity of lipid structures and pathways involved in the formation of membrane lipids when compared to eukaryotes. When studying the diversity of bacterial membrane lipids, the ultimate goal probably should be to have a complete catalog of the lipids that are formed when bacteria are growing in their natural habitats and understand their functions within the physiology of the organisms.
Although we have come a long way, there is still much work to do. What are the problems and what has to be done to fill these information gaps? Only a few bacteria can be easily grown in the laboratory in pure cultures.
Most bacteria of the environment still escape our attempts to grow them in the laboratory. This means that almost everything we know about bacterial membrane lipids comes from a relatively small set of easily culturable bacteria.
We can probably expect to find several unknown lipid structures and unknown synthesis pathways in these unculturable bacteria.
One possibility to untap part of this variety of bacterial membrane lipids is to combine culturomics approaches with lipidomics Lagier et al. Culturomics is a systematic approach to find growth conditions of so-far unculturable bacteria. Once this or these conditions are defined, the organism's membrane lipid composition can be studied.
Examples of recently discovered lipid structures include a trimethylated OL in Planctomycetes , diacylserinophospholipids from Verrucomicrobia and diol-based lipids in thermophilic bacteria Moore et al. Currently, we have a basic knowledge about the lipid composition of major model bacteria and many of the genes and enzymes involved in their synthesis.
However, even the well-studied E. For decades lipids biochemists had the idea that not much new was to be discovered in E. Even in well-studied organisms such as E. Almost all CDP-alcohol phosphotransferases characterized play a role in phospholipid synthesis, the only known exceptions being a few enzymes involved in the synthesis of inositol-based compatible solute in hyperthermophilic bacteria and archaea Brito et al.
It can be expected that many new structures and activities will be identified. Bacteria are cultured in the laboratory under conditions that in most cases do not reflect the natural habitat. The membrane lipid composition of bacteria is not a constant but is responding to the environmental conditions. If we want to understand what role the membrane lipids play for the functioning of a bacterium but also during the interaction with other bacteria or eukaryotes, it is essential to study the bacteria within their habitat.
Experimental approaches should be developed and improved that allow the study of bacterial membrane lipids in the natural environment or try to take the habitat to the laboratory. That means go to the sites where the bacteria are growing, sample and if possible study in situ or bring part of the habitat to the laboratory and study it there. One possibility is to combine the study of environmental lipid samples as for example geobiochemists do when analyzing samples from marine sediments, with metagenomic studies.
The metagenomic study will inform us what bacteria are present in the habitat and the lipid analysis will show what membrane lipids are present. In this model metadata such as the environmental conditions can be included, so that a modeling should be possible.
Finally, the perfect experimental approach would be to reconstitute the natural environment in the lab and use site-directed mutants in this experimental setup to follow how changes or conditions affect membrane composition and interactions between organisms. Sulfonolipids are molecular determinants of gliding motility Nature 9. Google Scholar. Membrane lipids in Agrobacterium tumefaciens : biosynthetic pathways and importance for pathogenesis Front Plant Sci 5 Limisphaera ngatamarikiensis gen.
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