The Heterokonta or Stramenopile phylum comprises clades of unicellular photosynthetic species, which are promising for a broad range of biotechnological applications, based on their capacity to capture atmospheric CO2 via photosynthesis and produce biomolecules of interest. These molecules include triacylglycerol (TAG) loaded inside specific cytosolic bodies, called the lipid droplets (LDs). Understanding TAG production and LD biogenesis and function in photosynthetic stramenopiles is therefore essential, and is mostly based on the study of a few emerging models, such as the pennate diatom Phaeodactylum tricornutum and eustigmatophytes, such as Nannochloropsis and Microchloropsis species. The biogenesis of cytosolic LD usually occurs at the level of the endoplasmic reticulum. However, stramenopile cells contain a complex plastid deriving from a secondary endosymbiosis, limited by four membranes, the outermost one being connected to the endomembrane system. Recent cell imaging and proteomic studies suggest that at least some cytosolic LDs might be associated to the surface of the complex plastid, via still uncharacterized contact sites. The carbon length and number of double bonds of the acyl groups contained in the TAG molecules depend on their origin. De novo synthesis produces long-chain saturated or monounsaturated fatty acids (SFA, MUFA), whereas subsequent maturation processes lead to very long-chain polyunsaturated FA (VLC-PUFA). TAG composition in SFA, MUFA, and VLC-PUFA reflects therefore the metabolic context that gave rise to the formation of the LD, either via an early partitioning of carbon following FA de novo synthesis and/or a recycling of FA from membrane lipids, e.g., plastid galactolipids or endomembrane phosphor- or betaine lipids. In this review, we address the relationship between cytosolic LDs and the complex membrane compartmentalization within stramenopile cells, the metabolic routes leading to TAG accumulation, and the physiological conditions that trigger LD production, in response to various environmental factors.
Appearance of oxygenic photosynthesis in Cyanobacteria is a major event in the evolution of Life. It had an irreversible impact on our planet, promoting the Great Oxygenation Event (GOE), ~2.4 b.y.a. Ancient Cyanobacteria predating the GOE were Gloeobacter-type cells, having no thylakoids. They hosted photosystems in their cytoplasmic membrane. The driver of the GOE was proposed to be the transition from unicellular to filamentous Cyanobacteria. However, the appearance of thylakoids expanded the photosynthetic surface by multiple logs: this multiplier effect would be more coherent with an impact on the atmosphere. Primitive thylakoids self-organize as concentric parietal uninterrupted multilayers. The quest for their origin resists vesicular-based scenarios. This review reports studies supporting that Hexagonal II-forming gluco- and galactolipids at the periphery of the cytosolic membrane could be turned within nanoseconds and without any external source of energy into membrane multilayers. Comparison of lipid biosynthetic pathways further shows that ancient Cyanobacteria contained only one anionic Lamellar-forming lipid, phosphatidylglycerol. Acquisition of sulfoquinovosyldiacylglycerol biosynthesis correlates with thylakoid emergence, possibly enabling a sufficient provision of anionic lipids to trigger an Hexagonal II-to-Lamellar phase transition. With this non-vesicular lipid-phase transition, a framework is also available to reexamine the role of companion proteins in thylakoid biogenesis processes.
Organic carbon fixed through the Calvin Cycle can be diverted towards different metabolic fates within and beyond the plastids of photosynthetic eukaryotes. These include export to the cytoplasm and mitochondrial respiration; gluconeogenesis of storage compounds; and the anabolic synthesis of lipids, amino acids and cofactors via the plastidial pyruvate hub. In plants, pyruvate is principally synthesised via the lower half of glycolysis-gluconeogenesis in the cytoplasm, although a secondary plastid-targeted pathway in non-photosynthetic tissue directly links glyceraldehyde-3-phosphate to the pyruvate hub. Here, we characterize a complete plastidial lower half glycolytic-gluconeogenic pathway in the photosynthetic plastids of diatoms, obligately photosynthetic eukaryotic algae that are important contributors to marine primary production. We show that the two enzymes required to complete plastidial glycolysis-gluconeogenesis, plastidial Enolase and PGAM (bis-phospho-glycerate mutase), originated through recent duplications of mitochondria-targeted respiratory glycolytic isoforms. Through CRISPR-Cas9 mutagenesis and integrative omic analyses in the diatom Phaeodactylum tricornutum, we present evidence that this pathway functions to divert excess plastidial glyceraldehyde-3-phosphate into diverse fates accessed from the pyruvate hub, and may potentially also function in the gluconeogenic direction to permit more efficient management of cellular carbon. Considering meta-genomic data, we show that this pathway is of greater importance in polar and sub-polar oceans, in which diatoms dominate primary production; and considering experimental data, we show that this principally relates to the elongated photoperiods present at high latitudes. Our data provide insights into the functions of a poorly understood yet evolutionarily recurrent plastidial metabolic pathway, and a further explanation for the success of diatoms in the contemporary ocean.
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