Micromonas
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''Micromonas'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
in the family ''
Mamiellaceae IMamiellaceae is a family of green algae in the order Mamiellales.See the NCBI The National Center for Biotechnology Information (NCBI) is part of the United States National Library of Medicine (NLM), a branch of the National Institutes of H ...
''. Before the characterization in 2016 of a second species, ''Micromonas commoda'', ''Micromonas pusilla'' was considered to be the only species in the genus, which led to a disproportionate amount of research discussing a single species within the genus. It is suggested to be the dominant photosynthetic
picoeukaryote Picoeukaryotes are picoplanktonic eukaryotic organisms 3.0 µm or less in size. They are distributed throughout the world's marine and freshwater ecosystems and constitute a significant contribution to autotrophic communities. Though the ...
in some marine ecosystems. Unlike many marine algae, it is distributed widely in both warm and cold waters.Genomes of Two Strains of Micromonas Algae Show Surprising Diversity
, ''Alternative Energy Newswire'', April 10, 2009
It is a strong swimmer and exhibits a phototaxic response. ''Micromonas pusilla'' is divided into 3 to 5 different
clades A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, t ...
despite their similarity in morphologies and habitats. Varying ratios of clades contribute to the ''M. pusilla'' population throughout the marine ecosystem leading to the hypothesis of clades arising based on niche occupation and susceptibility to virus infection.


Discovery

''Micromonas pusilla'' is considered the first
picoplankton Picoplankton is the fraction of plankton composed by cells between 0.2 and 2 μm that can be either prokaryotic and eukaryotic phototrophs and heterotrophs: * photosynthetic * heterotrophic They are prevalent amongst microbial plankton communit ...
studied, when it was discovered and named ''Chromulina pusilla'' in the 1950s by R. Butcher. Later, electron micrographs by the English scientists, Irene Manton and Mary Park, in the 1960s provided further details on ''M. pusilla''.


Cell morphology and structure

''Micromonas'' is a group of small
unicellular A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
pear-shaped micro-algae that do not have a
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
. Just like other members in the class, they have a single
mitochondrion A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
and a single
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
, which covers almost half of the cell. They are able to swim due to the presence of a scale-less
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
. The axonemal structure of the
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
for this genus is different in that the peripheral microtubules do not extend up to the termination of the central pair of
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
s, allowing a visible investigation of the motion of the central pair. In ''Micromonas'', the central pair constantly rotates in an anti-clockwise direction, despite the motion of other components of the
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
. While the cell size, shape and the location of insertion of the
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
into the cell are similar among strains and genetic clades, the variation in respective hair point length results in different lengths of the
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
within the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
.


Antibiotic

The
antibiotic susceptibility Antibiotic sensitivity testing or antibiotic susceptibility testing is the measurement of the susceptibility of bacteria to antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important typ ...
was determined using a single strain of ''M. pusilla'' with the purpose to produce
axenic In biology, axenic (, ) describes the state of a culture in which only a single species, variety, or strain of organism is present and entirely free of all other contaminating organisms. The earliest axenic cultures were of bacteria or unicellul ...
cultures to be used in studies and experiments. The strain of ''M.pusilla'' was tested with a range of antibiotics to determine the possible effects of the particular antibiotic. Resistance:
benzylpenicillin Benzylpenicillin, also known as penicillin G (PenG) or BENPEN, and in military slang "Peanut Butter Shot" is an antibiotic used to treat a number of bacterial infections. This includes pneumonia, strep throat, syphilis, necrotizing enterocolitis ...
,
gentamicin Gentamicin is an antibiotic used to treat several types of bacterial infections. This may include bone infections, endocarditis, pelvic inflammatory disease, meningitis, pneumonia, urinary tract infections, and sepsis among others. It is not e ...
,
kanamycin Kanamycin A, often referred to simply as kanamycin, is an antibiotic used to treat severe bacterial infections and tuberculosis. It is not a first line treatment. It is used by mouth, injection into a vein, or injection into a muscle. Kanamycin ...
,
neomycin Neomycin is an aminoglycoside antibiotic that displays bactericidal activity against gram-negative aerobic bacilli and some anaerobic bacilli where resistance has not yet arisen. It is generally not effective against gram-positive bacilli and an ...
,
streptomycin Streptomycin is an antibiotic medication used to treat a number of bacterial infections, including tuberculosis, ''Mycobacterium avium'' complex, endocarditis, brucellosis, ''Burkholderia'' infection, plague, tularemia, and rat bite fever. Fo ...
Sensitive:
chloramphenicol Chloramphenicol is an antibiotic useful for the treatment of a number of bacterial infections. This includes use as an eye ointment to treat conjunctivitis. By mouth or by injection into a vein, it is used to treat meningitis, plague, cholera, a ...
,
polymyxin B Polymyxin B, sold under the brand name Poly-Rx among others, is an antibiotic used to treat meningitis, pneumonia, sepsis, and urinary tract infections. While it is useful for many Gram negative infections, it is not useful for Gram positive infe ...
For ''M. pusilla'', sensitivity towards an antibiotic is likely defined by the impairment of growth, rather than a lethal effect, when subjected to
bactericidal A bactericide or bacteriocide, sometimes abbreviated Bcidal, is a substance which kills bacteria. Bactericides are disinfectants, antiseptics, or antibiotics. However, material surfaces can also have bactericidal properties based solely on their ...
levels of that particular antibiotic. The susceptibility of other strains of ''M. pusilla'' towards this set of antibiotics should be the same.


Genetics


Evolutionary history

''Micromonas'' diverged early on from the lineage that led to all modern terrestrial plants. Individual species have very similar
18S ribosomal RNA 18S ribosomal RNA (abbreviated 18S rRNA) is a part of the ribosomal RNA. The S in 18S represents Svedberg units. 18S rRNA is an SSU rRNA, a component of the eukaryotic ribosomal small subunit (40S). 18S rRNA is the structural RNA for the small c ...
gene sequences, a comparison often used to determine microscopic speciation, however, only 90% of different genes are shared between all ''Micromonas'' species. This presence or absence of complete genes, compared with only small changes of sequence to specific genes, suggests that ''Micromonas'' is the result of intensive
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between Unicellular organism, unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offsprin ...
.


Strain Isolation

The original ''Micromonas'' reference
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
was created from a
strain Strain may refer to: Science and technology * Strain (biology), variants of plants, viruses or bacteria; or an inbred animal used for experimental purposes * Strain (chemistry), a chemical stress of a molecule * Strain (injury), an injury to a mu ...
, RCC299, first isolated in 1998 from an Equatorial Pacific sample. This strain has been continuously cultured for two decades and is available from the Roscoff Culture Collection. In 2005, a monoclonal culture of the strain was isolated. The axenic strain is available from the Center for Culture of Marine Phytoplankton, under the name CCMP2709. Currently, a separate stain that was isolated from temperate coastal waters is being sequenced.


Genome Structure

The entire ''Micromonas sp.'' genome was first shotgun sequenced in 2014. ''Micromonas'' has about 19Mb, however this varies slightly between species and strain. This is made up of 17 chromosomes and contains 59% GC content. Based on
pen reading frames A pen is a common writing instrument that applies ink to a surface, usually paper, for writing or drawing. Early pens such as reed pens, quill pens, dip pens and ruling pens held a small amount of ink on a nib or in a small void or cavity whic ...
the genome codes for about 10000 proteins and 70 functional RNAs.


Cellular mechanisms


Cell growth and division

''Micromonas'' reproduces asexually through fission. It has been observed that ''M. pusilla'' shows variability in optical characteristics, for example cell size and light scattering, throughout the day. There is an increase in these measurements during the period with light, followed by a decrease during period without light. This coincides with the findings that
proteomic Proteomics is the large-scale study of proteins. Proteins are vital parts of living organisms, with many functions such as the formation of structural fibers of muscle tissue, enzymatic digestion of food, or synthesis and replication of DNA. In ...
profiles change over the diel cycle, with an increase in expression of proteins related to
cell proliferation Cell proliferation is the process by which ''a cell grows and divides to produce two daughter cells''. Cell proliferation leads to an exponential increase in cell number and is therefore a rapid mechanism of tissue growth. Cell proliferation re ...
,
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
and
cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
restructuring in the dark when cells start dividing and become smaller. However, the expression levels of genes and proteins can still vary within the same
metabolic pathway In biochemistry, a metabolic pathway is a linked series of chemical reactions occurring within a cell. The reactants, products, and intermediates of an enzymatic reaction are known as metabolites, which are modified by a sequence of chemical reac ...
. It has also been suggested that the structure of 3’ UTR may play a role in the regulatory system.


Light-harvesting system

''Micromonas'' species still share the same collection of
photosynthetic pigment A photosynthetic pigment (accessory pigment; chloroplast pigment; antenna pigment) is a pigment that is present in chloroplasts or photosynthetic bacteria and captures the light energy necessary for photosynthesis. List of photosynthetic pigmen ...
s as the members of the class ''
Mamiellophyceae Mamiellophyceae is a class of green algae in the division Chlorophyta Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the u ...
,'' which includes the common pigments chlorophyll a and
chlorophyll b } Chlorophyll ''b'' is a form of chlorophyll. Chlorophyll ''b'' helps in photosynthesis by absorbing light energy. It is more soluble than chlorophyll ''a'' in polar solvents because of its carbonyl group. Its color is green, and it primarily ...
, as well as prasinoxanthin (xanthophyll K), the first algal
carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic compound, organic pigments that are produced by plants and algae, as well as several bacteria, and Fungus, fungi. Carotenoids give the characteristic color to pumpki ...
being assigned with a structure that has a γ-end group. It has been discovered that most of its
xanthophyll Xanthophylls (originally phylloxanthins) are yellow pigments that occur widely in nature and form one of two major divisions of the carotenoid group; the other division is formed by the carotenes. The name is from Greek (, "yellow") and (, "l ...
s are in the oxidized state and show similarities to ones possessed by other important marine
plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
s like diatoms,
golden Golden means made of, or relating to gold. Golden may also refer to: Places United Kingdom *Golden, in the parish of Probus, Cornwall * Golden Cap, Dorset *Golden Square, Soho, London *Golden Valley, a valley on the River Frome in Gloucestershi ...
and
brown algae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and po ...
, and
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s. In addition, there is another pigment called Chl cCS-170 can be found in some strains of ''Micromonas'' and ''
Ostreococcus ''Ostreococcus'' is a genus of unicellular coccoid or spherically shaped green algae belonging to the class Mamiellophyceae. It includes prominent members of the global picoplankton community, which plays a central role in the oceanic carbon cyc ...
'' living in deeper part of the ocean, which may indicate a potential adaptation for organisms that reside under low light intensity. The light-harvesting complexes of ''Micromonas'' are distinguishable from other
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
in terms of pigment composition and stability under unfavorable conditions. It has been showed that these proteins use three different pigments for light harvesting, and they are resistant to high temperature and the presence of detergent.


Peptidoglycan biosynthesis

Even though the
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s, which are suggested to be originated from
Cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
via
endosymbiosis An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
, from ''Micromonas'' do not have a surrounding
peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...
layer, the
peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...
biosynthesis pathway is found to be complete in ''M.'' ''pusilla'' and partial in ''M. commoda'', with the presence of some relevant enzymes only. While the role of this pathway for ''Micromonas'' is still under investigation, this observation shows a lineage for different species of ''Micromonas'' along with glaucophyte algae which still have their
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s covered with
peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...
.


Ecological significance

''Micromonas'' make up a significant amount of picoplanktonic biomass and productivity in both oceanic and coastal regions. The abundance of ''Micromonas'' has increased over the past decade. Evidence shows these spikes in numbers are induced through climate change, which has been felt more drastically in the Arctic. In past years it was thought that green algal species were solely photosynthetic only to discover that was not the case. ''Micromonas'', as well as the Prasinophytes, have taken on mixotrophic lifestyles and have huge impacts on prokaryotic populations within the Arctic. Due to the large consumption of prokaryotes by ''Micromonas'', studies are suggesting photosynthetic picoeukaryotes will soon dominate the primary productivity and bacterivory of Arctic systems. Laboratory studies have shown that species within a single genus have altered their mixotrophic strategies to their distinct environments. These environments can differ through light intensity, nutrient availability and size of prey which all have shown clade-specific alterations to maximize efficiencies.


Viral infection

Virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsky's 1 ...
es are important in the balance of marine ecosystem by regulating the composition of microbial communities, but their behaviors can be affected by several factors including temperature, mode of infection and host conditions. There is an increasing number of ''Micromonas''-infecting virus being discovered and studied.


''Micromonas pusilla'' virus

There are currently 45 viral strains identified that coexist with ''M. pusilla'' populations. Virus infectivity is dependent on the host strain, light availability and virus adsorption. Per day average of death due to virus lysis is estimated to be about 2 to 10% of the ''M. pusilla'' population. * ''Micromonas pusilla'' reovirus (MpRV): The first isolation of a
reovirus ''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
that infects protist. This virus is found to be bigger than other members of the family.


''Micromonas polaris'' virus

It is the first
phycodnavirus ''Phycodnaviridae'' is a family of large (100–560 kb) double-stranded DNA viruses that infect marine or freshwater eukaryotic algae. Viruses within this family have a similar morphology, with an icosahedral capsid (polyhedron with 20 fac ...
being isolated from polar ocean waters. It can infect ''M. polaris'', which is the polar ecotype of ''Micromonas'' that has adapted to waters with low temperatures. Evidence suggests that the increase in temperature due to climate change may shift the clonal composition of both the virus and host.


Metabolic engineering

With the growing population in the world, there is an increased demand for wild fishes and algae for their source of
polyunsaturated fatty acids Polyunsaturated fatty acids (PUFAs) are fatty acids that contain more than one double bond in their backbone. This class includes many important compounds, such as essential fatty acids and those that give drying oils their characteristic proper ...
(PUFA), which is required for growth and development, as well as the maintenance of health in humans. Recent research is investigating an alternative mechanism for production of PUFA by using acyl-CoA Δ6-desaturase, an enzyme present in ''M. pusilla'', with plants. The ''M. pusilla'' strain of acyl-CoA Δ6-desaturase is highly effective in the polyunsaturated fatty acid synthesis pathway due to its strong binding preference for omega-3 substrates in land plants.


References


External links


Genes from tiny algae shed light on big role managing carbon in world's oceans
{{Taxonbar, from=Q6839686 Chlorophyta genera Mamiellophyceae Marine microorganisms