Nannochloropsis

Last updated

Nannochloropsis
15 3klein2.jpg
Nannochloropsis viewed under a light microscope
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Stramenopiles
Phylum: Gyrista
Subphylum: Ochrophytina
Class: Eustigmatophyceae
Order: Eustigmatales
Family: Monodopsidaceae
Genus: Nannochloropsis
D.J.Hibberd, 1981 [1]
Type species
Nannochloropsis oculata
(Droop 1955) Hibberd 1981

Nannochloropsis is a genus of algae comprising six known species. The genus in the current taxonomic classification was first termed by Hibberd (1981). [2] The species have mostly been known from the marine environment but also occur in fresh and brackish water. [3] All of the species are small, nonmotile spheres which do not express any distinct morphological features that can be distinguished by either light or electron microscopy. The characterisation is mostly done by rbcL gene and 18S rRNA sequence analysis. [4]

Contents

The algae of the genus Nannochloropsis differ from other related microalgae in that they have chlorophyll a and completely lack chlorophyll b and chlorophyll c . In addition they are able to build up a high concentrations of a range of pigments such as astaxanthin, zeaxanthin and canthaxanthin. [5] They have a diameter of about 2 to 3 micrometers [6] and a very simple ultrastructure with reduced structural elements compared to neighbouring taxa.

Nannochloropsis is considered a promising alga for industrial applications because of its ability to accumulate high levels of polyunsaturated fatty acids. [7] [8] Moreover, it shows promising features that can allow genetic manipulation aimed at the genetic improvement of the current oleaginous strains. Various species of Nannochloropsis indeed are transfectable and there has been evidence that some strains are able to perform homologous recombination. [9] At the moment it is mainly used as an energy-rich food source for fish larvae and rotifers. Nevertheless, it has raised growing interest also for the investigation of biofuel production from photosynthetic organisms. (see Nannochloropsis and biofuels). Nannochloropsis is actually in use as food additive for human nutrition and it is also served at Restaurant "A Poniente" of El Puerto de Santa María (Cádiz, Spain) close to the natural environment where Nannochloropsis gaditana was first isolated and still grows. [10] A 2020 study suggests it could be used for a highly performant, sustainable fish-free feed for farmed fish. [11] [12]

Species

The species currently recognised are:

Sequenced genomes

The scientific community has obtained the genomic sequence of different strains of Nannochloropsis belonging to two species: N. gaditana [14] [15] and N. oceanica. [16] [17] A genome portal based on the N. gaditana B-31 genome allows accessing much of the genomic information that concerns this micro-organism, moreover dedicated web pages are also available for the genomes of N. gaditana CCMP526 and N. oceanica CCMP1779.

The genomes of the sequenced Nannochloropsis strains were between 28.5 and 29 Mega bases long, they had high density of genes, reduced intron content, short intergenic regions [14] and very limited presence of repetitive sequences. [16] The genes of the two species share extended similarity.

The analysis of the genomes revealed that these microalgae have set of genes for the synthesis and incorporation in the cell wall of cellulose and sulfated fucans and that they are able to store carbon in polymers of β-1,3- and β-1,6-linked glucose called chrysolaminarin. [14] An NMR analysis of whole algal cells which were cultivated in autotrophic growth [18] reports evidence of the presence of cellulose in the cell wall and of mobile chrysolaminarin, probably accumulated in solution in vacuoles inside the cell.

Comparison between the lipid metabolic genes of N. gaditana and of red/green/brown algae and diatoms provided some insights into the exemplary lipid production of Nannochloropsis cultures. The comparisons indeed highlighted the presence of an expanded repertoire of some of the genes involved in TAG assembly in Nannochloropsis. [15] Numerous TAG lipases, which can affect TAG metabolism through either TAG degradation or lipid remodeling, were identified in Nannochloropsis, [16] many of them belonging to a gene family which seems to be exclusive of Nannochloropsis. [19]

Analysis of the genomic data suggested the presence in both the two species of Nannochloropsis of regulatory RNA and in particular miRNA. [14] [16]

Various orthologs of known blue light sensing proteins were found in the genomes of Nannochloropsis suggesting possible circadian regulation. [14] [16]

A research community from Qingdao Institute of Bioenergy and Bioprocess Technology (QIBEBT ), Chinese Academy of Sciences, sequenced and compared six Nannochloropsis genomes that include two N. oceanica strains (IMET1 and CCMP531) and one strain from each of four other recognized species: N. salina (CCMP537), N. gaditana (CCMP526, which was previously reported [15] ), N. oculata (CCMP525) and N. granulata (CCMP529). [20] They found that the six genomes share key oleaginous traits, such as the gene dose expansion of selected lipid biosynthesis genes compared to green algae Chlamydomonas. The most prominent example of gene dose expansion is Diacylglycerol acyltransferase (DGAT), which catalyzes the last step of triacylglycerol (TAG) synthesis. There are 13 DGAT genes in Nannochloropsis, representing the highest gene dose in known genomes.

Through a comprehensive phylogenetic analysis, researchers proposed that among the 11 DGAT-2s, one gene might originate from the red algae related secondary endosymbiont, four from green algae related endosymbiont, and the other six from the eukaryotic host genome. In addition, a large proportion (15.3%) of TAG biosynthesis related genes were acquired by Nannochloropsis via horizontal gene transfer (HGT) from bacteria. Therefore, multiple genome pooling and horizontal genetic exchange, together with selective inheritance of lipid synthesis genes and species-specific gene loss, have led to the enormous genetic apparatus for oleaginousness and the wide genomic divergence among present-day Nannochloropsis spp.

Accumulation of oil in nitrogen deprivation

When cultured in normal growth conditions in f/2 medium, a standard algal culture growth medium, supplemented with nitrogen, Nannochloropsis cells have an oil content of about 30% of their dry weight. This oil can be used to produce biodiesel. In order to produce biodiesel, the oil content of Nannochloropsis cultures must be increased. Fatty acid biosynthesis in N. oceanica was shown to be integrated with the regulation of the cholesterol biosynthetic pathway, suggesting new genetic engineering or chemical biology approaches for enhanced oil production in microalgae. [21] On the other hand, various culturing conditions were observed to increase the average oil content per cell, supporting however only slow growth rates of the cultures (see the related section Nannochloropsis and biofuels), and decreasing the overall productivity. Among these conditions, nitrogen deprivation has been one of the most vastly studied. Studies have examined the behaviour of the cultures in nitrogen stress in various culturing set-ups, as well as the physiological and molecular response of the cells to nitrogen deprivation.

Various strains of Nannochloropsis were shown to accumulate up to 60–70% of their overall biomass as lipids in nitrogen limitation. [8] [22] In this condition not only the amount of lipids changes, but also the composition. It was observed that triacylglycerols largely increase in nitrogen depletion while other lipids, mainly polar glycerolipids, free fatty acids and diacylglycerols, slightly decrease. The increase in triacylglycerols in nitrogen depletion is much larger than the decrease in the other glycerolipids, supporting the hypothesis that the cells actively synthesise new triacylglycerols rather than just transforming the existing lipids into triacylglycerols. [23]

While triacylglycerols accumulate and lipid droplets enlarge, other important morphological changes take place. The chloroplasts became smaller and the amount of lipids that are characteristic of the photosynthetic membranes decrease significantly. Photosynthetic yield decreases in the nitrogen depleted cells, but it is still sufficient to drive cell survival and growth. If photosynthesis is inhibited indeed, Nannochloropsis cells stop growing, indicating that they rely on photosynthetic solar energy conversion and not on stored carbohydrates or lipids as energy source. [23]

Nannochloropsis cultivated in normal growth condition and deprived of a nitrogen source continues growing for 4–5 days. The analysis of gene expression reveals that mechanisms for nitrogen assimilation and redistribution are activated in the cells in nitrogen scarcity and allow survival through a partial reorganisation of the cellular metabolism. The genes responsible for the incorporation of free ammonium into glutamate (L-glutamine + 2-oxoglutarate + NADPH + H+ ←→ 2 L-glutamate + NADP+) are up-regulated . Their up-regulation increases the capability of the cells to assimilate minimal amount of ammonium eventually present in the medium and, more importantly, allows them to assimilate the intracellular ammonium that can be released by degradative processes. Transcriptomic data of nitrogen-deprived Nannochloropsis cultures show that various enzymes responsible for degradative processes that release ammonium are indeed up regulated. Moreover, the expression of genes coding for proteins involved in controlled degradation of proteins increase, together with others involved in the formation of cytosolic sequestering vesicles used for degradation and recycling of cellular components. [14]

The machinery of gene expression of chloroplast and mitochondrion is down regulated in Nannochloropsis in response to nitrogen depletion. [14] [15]

Corteggiani Carpinelli and coworkers (2013) [14] report a complete analysis of the gene expression of Nannochloropsis cultures grown in normal conditions and nitrogen deprivation for 3 days and 6 days. Data on similar conditions were also collected by Radakovits et al. (2012) [15] and Vieler et al. (2012). [16] All of this data show that genes involved in fatty acid and triacylglycerol biosynthesis are always abundant in the cells and their expression is not correlated with the amount of oil accumulated. Also the expression of the genes involved in triacylglycerols degradation is not significantly down-regulated as triacylglycerols accumulate. The general conclusion suggested by the experimental data is that Nannochloropsis constitutively produces triacylglycerols and that the metabolic reorganisation that follows nitrogen deprivation increases the flux of substrates through this pathway, which is in turn capable to sustain the increased metabolic flux. Corteggiani Carpinelli and coworkers (2013) [14] advance the hypothesis that, in their experimental conditions, photosynthesis is the main energy source and the down-regulation of the metabolic activity of the mitochondrion is determinant in increasing the amount of substrates that enter the fatty acid biosynthetic pathway. If more precursors are available, more fatty acids are synthesised and as a consequence more triacylglycerols are produced and accumulated into the cells as oil droplets.

Dong et al. (2013) [24] report a quantification of the abundance of various proteins in cultures of Nannochloropsis grown with a supplement of CO2 and deprived of nitrogen. Despite the different experimental conditions, also the proteomic study seems to support the hypothesis that the accumulation of triacylglycerols is due to an increase of the metabolic flux through the fatty acid biosynthetic pathway. The authors advance the hypothesis that, in their experimental conditions, the degradation of storage sugars and the up-regulation of glycolysis are responsible for the increase of substrates through the pathway.

More recently Li and coworkers (2014) [25] collected extensive experimental data from cultures of Nannochloropsis oculata IMET1 grown in nitrogen sufficient and nitrogen depleted media. According to their analysis it is the catabolism of carbohydrates and proteins together with the up-regulation of genes assigned to various pathways (the cytosolic glycolysis pathway, which produces pyruvate; the PDHC bypass, which yields additional acetyl-CoA; and the coupling of TCA reactions with mitochondrial β-oxidation) that have to be claimed for increasing the supply of carbon precursors to the fatty acid biosynthetic pathway. Nevertheless, the authors suggest that it isn't the increased synthesis of fatty acids that leads to accumulation of oil inside the cells, but it is rather the up-regulation of genes involved in the assembly of triacylglycerols that determines the final oil concentration. In other words, under nitrogen depletion, it is the up-regulation of the transcript level of the key genes in triacylglycerol assembly, rather than those in fatty acids biosynthesis, that leads to accelerated triacylglycerol production.

In 2017, a study jointly funded by ExxonMobil and Synthetic Genomics achieved a breakthrough in lipid percentages in Nannochloropsis gaditana through applications of the CRISPR-Cas9 reverse-genetics pipeline and nitrogen starving—improving partitioning of total carbon to lipids from 20% (wild type) to 40-45% (mutant type). [26]

See also

Related Research Articles

<span class="mw-page-title-main">Microalgae</span> Microscopic algae

Microalgae or microphytes are microscopic algae invisible to the naked eye. They are phytoplankton typically found in freshwater and marine systems, living in both the water column and sediment. They are unicellular species which exist individually, or in chains or groups. Depending on the species, their sizes can range from a few micrometers (μm) to a few hundred micrometers. Unlike higher plants, microalgae do not have roots, stems, or leaves. They are specially adapted to an environment dominated by viscous forces.

<span class="mw-page-title-main">Eicosapentaenoic acid</span> Chemical compound

Eicosapentaenoic acid is an omega-3 fatty acid. In physiological literature, it is given the name 20:5(n-3). It also has the trivial name timnodonic acid. In chemical structure, EPA is a carboxylic acid with a 20-carbon chain and five cis double bonds; the first double bond is located at the third carbon from the omega end.

<span class="mw-page-title-main">Algaculture</span> Aquaculture involving the farming of algae

Algaculture is a form of aquaculture involving the farming of species of algae.

The Aquatic Species Program was a research program in the United States launched in 1978 by President Jimmy Carter and was funded by the United States Department of Energy, which over the course of nearly two decades looked into the production of energy using algae. Initially, the funding of the Aquatic Species Program was to develop renewable fuel for transportation. Later, the program focused on producing bio-diesel from algae. The research program was discontinued in 1996. The research staff compiled their work and conclusions into a 1998 report.

<i>Botryococcus braunii</i> Species of alga

Botryococcus braunii is a green, pyramid-shaped planktonic microalga that is of potentially great importance in the field of biotechnology. Colonies held together by a lipid biofilm matrix can be found in temperate or tropical oligotrophic lakes and estuaries, and will bloom when in the presence of elevated levels of dissolved inorganic phosphorus. The species is notable for its ability to produce high amounts of hydrocarbons, especially oils in the form of Triterpenes, that are typically around 30–40% of their dry weight. Compared to other green alge species it has a relatively thick cell wall that is accumulated from previous cellular divisions; making extraction of cytoplasmic components rather difficult. Much of the useful hydrocarbon oil is outside of the cell.

<i>Scenedesmus</i> Genus of green algae

Scenedesmus is a genus of green algae, in the class Chlorophyceae. They are colonial and non-motile. They are one of the most common components of phytoplankton in freshwater habitats worldwide.

Auxenochlorella protothecoides, formerly known as Chlorella protothecoides, is a facultative heterotrophic green alga in the family Chlorellaceae. It is known for its potential application in biofuel production. It was first characterized as a distinct algal species in 1965, and has since been regarded as a separate genus from Chlorella due its need for thiamine for growth. Auxenochlorella species have been found in a wide variety of environments from acidic volcanic soil in Italy to the sap of poplar trees in the forests of Germany. Its use in industrial processes has been studied, as the high lipid content of the alga during heterotrophic growth is promising for biodiesel; its use in wastewater treatment has been investigated, as well.

<i>Choricystis</i> Genus of algae

Choricystis is a genus of green algae in the class Trebouxiophyceae, considered a characteristic picophytoplankton in freshwater ecosystems. Choricystis, especially the type species Choricystis minor, has been proposed as an effective source of fatty acids for biofuels. Choricystis algacultures have been shown to survive on wastewater. In particular, Choricystis has been proposed as a biological water treatment system for industrial waste produced by the processing of dairy goods.

<span class="mw-page-title-main">Algae fuel</span> Use of algae as a source of energy-rich oils

Algae fuel, algal biofuel, or algal oil is an alternative to liquid fossil fuels that uses algae as its source of energy-rich oils. Also, algae fuels are an alternative to commonly known biofuel sources, such as corn and sugarcane. When made from seaweed (macroalgae) it can be known as seaweed fuel or seaweed oil.

In enzymology, a long-chain-alcohol O-fatty-acyltransferase is an enzyme that catalyzes the chemical reaction

<i>Nannochloropsis</i> and biofuels

Nannochloropsis is a genus of alga within the heterokont line of eukaryotes, that is being investigated for biofuel production. One marine Nannochloropsis species has been shown to be suitable for algal biofuel production due to its ease of growth and high oil content, mainly unsaturated fatty acids and a significant percentage of palmitic acid. It also contains enough unsaturated fatty acid linolenic acid and polyunsaturated acid for a quality biodiesel.

<span class="mw-page-title-main">Culture of microalgae in hatcheries</span>

Microalgae or microscopic algae grow in either marine or freshwater systems. They are primary producers in the oceans that convert water and carbon dioxide to biomass and oxygen in the presence of sunlight.

Single cell oil, also known as Microbial oil consists of the intracellular storage lipids, triacyglycerols. It is similar to vegetable oil, another biologically produced oil. They are produced by oleaginous microorganisms, which is the term for those bacteria, molds, algae and yeast, which can accumulate 20% to 80% lipids of their biomass. The accumulation of lipids take place by the end of logarithmic phase and continues during station phase until carbon source begins to reduce with nutrition limitation.

Schizochytrium is a genus of unicellular eukaryotes in the family Thraustochytriaceae, which are found in coastal marine habitats. They are assigned to the Stramenopiles (heterokonts), a group which also contains kelp and various microalgae.

Sammy Boussiba is a professor emeritus at the French Associates Institute for Agriculture and Biotechnology of Drylands at the Jacob Blaustein Institutes for Desert Research at Ben-Gurion University of the Negev, Israel.

Monoraphidium neglectum is a single-cell green alga of the family Selenastraceae. Cells are free-floating (planktonic), fusiform and sometimes arched in outline, with rounded-pointed tips. They are 16–30 μm long by 3–4.5 μm wide. The chloroplast is single per cell and lacks a pyrenoid. It reproduces by forming two, four or eight autospores.

The MSU-DOE Plant Research Laboratory (PRL), commonly referred to as Plant Research Lab, is a research institute funded to a large extent by the U.S. Department of Energy Office of Science and located at Michigan State University (MSU) in East Lansing, Michigan. The Plant Research Lab was founded in 1965, and it currently includes twelve laboratories that conduct collaborative basic research into the biology of diverse photosynthetic organisms, including plants, bacteria, and algae, in addition to developing new technologies towards addressing energy and food challenges.

<span class="mw-page-title-main">Thraustochytrids</span> Order of eukaryotes

Thraustochytrids are single-celled saprotrophic eukaryotes (decomposers) that are widely distributed in marine ecosystems, and which secrete enzymes including, but not limited to amylases, proteases, phosphatases. They are most abundant in regions with high amounts of detritus and decaying plant material. They play an important ecological role in mangroves, where they aid in nutrient cycling by decomposing decaying matter. Additionally, they contribute significantly to the synthesis of omega-3 polyunsaturated fatty acids (PUFAs): docosahexaenoic acid (DHA), and eicosapentaenoic acid (EPA), which are essential fatty acids for the growth and reproduction of crustaceans. Thraustochytrids are members of the class Labyrinthulea, a group of protists that had previously been incorrectly categorized as fungi due to their similar appearance and lifestyle. With the advent of DNA sequencing technology, labyrinthulomycetes were appropriately placed with other stramenopiles and subsequently categorized as a group of Labyrinthulomycetes.

Christoph Benning is a German–American plant biologist. He is an MSU Foundation Professor and University Distinguished Professor at Michigan State University. Benning's research into lipid metabolism in plants, algae and photosynthetic bacteria, led him to be named Editor-in-Chief of The Plant Journal in October 2008.

An oleaginous microorganism is a type of microbe that accumulates lipid as a normal part of its metabolism. Oleaginous microbes may accumulate an array of different lipid compounds, including polyhydroxyalkanoates, triacylglycerols, and wax esters. Various microorganisms, including bacteria, fungi, and yeast, are known to accumulate lipids. These organisms are often researched for their potential use in producing fuels from waste products.

References

  1. ↑ Hibberd, D. J. (1981). "Notes on the taxonomy and nomenclature of the algal classes Eustigmatophyceae and Tribophyceae (synonym Xanthophyceae)". Botanical Journal of the Linnean Society. 82 (2): 93–119. doi:10.1111/j.1095-8339.1981.tb00954.x.
  2. ↑ Hibberd (1981). "Notes on the taxonomy and nomenclature of the algal classes Eustigmatophyceae and Tribophyceae (Synonym Xanthophyceae)". Botanical Journal of the Linnean Society. 82 (2): 93–119. doi:10.1111/j.1095-8339.1981.tb00954.x.
  3. ↑ Fawley (2007). "Observations on the Diversity and Ecology of Freshwater Nannochloropsis (Eustigmatophyceae), with Descriptions of New Taxa". Protist. 158 (3): 325–336. doi:10.1016/j.protis.2007.03.003. PMID   17576099.
  4. ↑ Andersen, R. A.; Brett, R. W.; et al. (1998). "Phylogeny of the Eustigmatophyceae based upon 18s rDNA, with emphasis on Nannochloropsis". Protist. 149 (1): 61–74. doi:10.1016/S1434-4610(98)70010-0. PMID   23196114.
  5. ↑ Lubian, L. M.; et al. (2000). "Nannochloropsis (Eustigmatophyceae) as source of commercially valuable pigments". Journal of Applied Phycology. 12 (3/5): 249–255. doi:10.1023/A:1008170915932. S2CID   620002.
  6. ↑ Kandilian, R.; Lee, E.; Pilon, L. (2013). "Radiation and optical properties of Nannochloropsis oculata grown under different irradiances and spectra". Bioresource Technology. 137: 63–73. doi:10.1016/j.biortech.2013.03.058. PMID   23587810.
  7. ↑ Assaf Sukenik, Y. C. T. B. (1989). "Regulation of fatty acid composition by irradiance level in the Eustigmatophyte Nannochloropsis sp". Journal of Phycology. 25 (4): 686–692. doi:10.1111/j.0022-3646.1989.00686.x. S2CID   84590502.
  8. 1 2 Boussiba, S.; Vonshak, A.; Cohen, Z.; Avissar, Y.; Richmond, A. (1987). "Lipid and biomass production by the halotolerant microalga Nannochloropsis salina". Biomass. 12: 37–47. doi:10.1016/0144-4565(87)90006-0.
  9. ↑ Kilian, O.; Benemann, C. S. E.; Niyogi, K. K.; Vick, B. (2011). "High-efficiency homologous recombination in the oil-producing alga Nannochloropsis sp". Proc. Natl. Acad. Sci. U.S.A. 108 (52): 21265–21269. doi: 10.1073/pnas.1105861108 . PMC   3248512 . PMID   22123974.
  10. ↑ Lubian, L. M. (1982). "Nannochloropsis gaditana sp. nov., a new Eustigmatophyceae marina strain". Lazaroa. 4: 287–293.
  11. ↑ "Research breakthrough achieves fish-free aquaculture feed that raises key standards". phys.org. Retrieved 9 December 2020.
  12. ↑ Sarker, Pallab K.; Kapuscinski, Anne R.; McKuin, Brandi; Fitzgerald, Devin S.; Nash, Hannah M.; Greenwood, Connor (12 November 2020). "Microalgae-blend tilapia feed eliminates fishmeal and fish oil, improves growth, and is cost viable". Scientific Reports . 10 (1): 19328. Bibcode:2020NatSR..1019328S. doi:10.1038/s41598-020-75289-x. ISSN   2045-2322. PMC   7665073 . PMID   33184333. CC-BY icon.svg Available under CC BY 4.0.
  13. ↑ Huesemann, M.; Williams, P.; Edmundson, Scott J.; Chen, P.; Kruk, R.; Cullinan, V.; Crowe, B.; Lundquist, T. (September 2017). "The laboratory environmental algae pond simulator (LEAPS) photobioreactor: Validation using outdoor pond cultures of Chlorella sorokiniana and Nannochloropsis salina". Algal Research. 26: 39–46. doi: 10.1016/j.algal.2017.06.017 . ISSN   2211-9264. OSTI   1581797.
  14. 1 2 3 4 5 6 7 8 9 Corteggiani Carpinelli, E.; et al. (2013). "Chromosome scale genome assembly and transcriptome profiling of Nannochloropsis gaditana in nitrogen depletion". Molecular Plant. 7 (2): 323–335. doi: 10.1093/mp/sst120 . PMID   23966634.
  15. 1 2 3 4 5 Radakovits, R.; et al. (2012). "Draft genome sequence and genetic transformation of the oleaginous alga Nannochloropis gaditana". Nat Commun. 3 (2): 686. Bibcode:2012NatCo...3..686R. doi:10.1038/ncomms1688. PMC   3293424 . PMID   22353717.
  16. 1 2 3 4 5 6 Vieler, A.; et al. (2012). "Genome, Functional Gene Annotation, and Nuclear Transformation of the Heterokont Oleaginous Alga Nannochloropsis oceanica CCMP1779". PLOS Genet. 8 (11): e1003064. doi: 10.1371/journal.pgen.1003064 . PMC   3499364 . PMID   23166516.
  17. ↑ Liang, C.; et al. (2012). "De Novo Sequencing and Global Transcriptome Analysis of Nannochloropsis sp. (Eustigmatophyceae) Following Nitrogen Starvation". Bioenerg. Res. 6 (2): 494–505. doi:10.1007/s12155-012-9269-0. S2CID   207395605.
  18. ↑ Arnold, A. A.; et al. (2014). "Identification of lipid and saccharide constituents of whole microalgal cells by 13C solid-state NMR". Biochimica et Biophysica Acta (BBA) - Biomembranes. 1848 (1 Pt B): 369–377. doi: 10.1016/j.bbamem.2014.07.017 . PMID   25065670.
  19. ↑ Elisa Corteggiani Carpinelli e Nicola Vitulo Unpublished data available through the Nannochloropsis gaditana genome portal Archived 2013-10-29 at the Wayback Machine
  20. ↑ Wang, Dongmei; Ning, Kang; Li, Jing; et, al.; Xu, Jian (January 2014). "Nannochloropsis Genomes Reveal Evolution of Microalgal Oleaginous Traits". PLOS Genetics. 10 (1): e1004094. doi: 10.1371/journal.pgen.1004094 . PMC   3886936 . PMID   24415958.
  21. ↑ Lu; et al. (2014). "Regulation of the cholesterol biosynthetic pathway and its integration with fatty acid biosynthesis in the oleaginous microalga Nannochloropsis oceanica". Biotechnology for Biofuels. 7: 81. doi: 10.1186/1754-6834-7-81 . PMC   4052811 . PMID   24920959.
  22. ↑ Rodolfi, L; Zittelli, Chini; et al. (2008). "Microalgae for oil: strain selection, induction of lipid synthesis and outdoor mass cultivation in a low-cost photobioreactor". Biotechnology and Bioengineering. 102 (1): 100–12. doi:10.1002/bit.22033. PMID   18683258. S2CID   28285869.
  23. 1 2 Simionato, Diana; Block, Maryse A.; Rocca, Nicoletta La; Jouhet, Juliette; Maréchal, Eric; Finazzi, Giovanni; Morosinotto, Tomas (2013). "The response of Nannochloropsis gaditana to nitrogen starvation includes de novo biosynthesis of triacylglycerols, a decrease of chloroplast galactolipids, and reorganisation of the photosynthetic apparatus". Eukaryotic Cell. 12 (5): 665–676. doi:10.1128/EC.00363-12. PMC   3647774 . PMID   23457191.
  24. ↑ Dong, HP; Williams, E; Wang, DZ; Xie, ZX; Hsia, RC; Jenck, A; Halden, R; Li, J; Chen, F; Place, AR (2013). "Responses of Nannochloropsis oceanica IMET1 to long-term nitrogen starvation and recovery". Plant Physiology. 162 (2): 1110–1126. doi:10.1104/pp.113.214320. PMC   3668043 . PMID   23637339.
  25. ↑ Li, Jing; Han, Danxiang; Wang, Dongmei; Ning, Kang; Jia, Jing; Wei, Li; Jing, Xiaoyan; Huang, Shi; Chen, Jie (2014). "Choreography of transcriptomes and lipidomes of Nannochloropsis reveals the mechanisms of oil synthesis in microalgae". Plant Cell. 26 (4): 1645–1665. doi:10.1105/tpc.113.121418. PMC   4036577 . PMID   24692423.
  26. ↑ Ajjawi, Imad; Verruto, John; Aqui, Moena; Soriaga, Leah B; Coppersmith, Jennifer; Kwok, Kathleen; Peach, Luke; Orchard, Elizabeth; Kalb, Ryan (2017). "Lipid production in Nannochloropsis gaditana is doubled by decreasing expression of a single transcriptional regulator". Nature Biotechnology. 35 (7): 647–652. doi:10.1038/nbt.3865. PMID   28628130. S2CID   8175045.
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.