Chlamydomonas reinhardtii

Chlamydomonas reinhardtii is a single-cell green alga about 10 micrometres in diameter that swims with two flagella. It has a cell wall made of hydroxyproline-rich glycoproteins, a large cup-shaped chloroplast, a large pyrenoid, and an eyespot that senses light.
"Chlamydomonas" species are widely distributed worldwide in soil and fresh water. "Chlamydomonas reinhardtii" is an especially well studied biological model organism, partly due to its ease of culturing and the ability to manipulate its genetics. When illuminated, "C. reinhardtii" can grow photoautotrophically, but it can also grow in the dark if supplied with organic carbon. Commercially, "C. reinhardtii" is of interest for producing biopharmaceuticals and biofuel, as well being a valuable research tool in making hydrogen.
History.
The "C. reinhardtii" wild-type laboratory strain c137 (mt+) originates from an isolate made near Amherst, Massachusetts, in 1945 by Gilbert M. Smith.
The species' name has been spelled several different ways because of different transliterations of the name from Russian: "reinhardi", "reinhardii", and "reinhardtii" all refer to the same species, "C. reinhardtii" Dangeard.
Model organism.
"Chlamydomonas" is used as a model organism for research on fundamental questions in cell and molecular biology such as:
There are many known mutants of "C. reinhardtii". These mutants are useful tools for studying a variety of biological processes, including flagellar motility, photosynthesis, and protein synthesis. Since "Chlamydomonas" species are normally haploid, the effects of mutations are seen immediately without further crosses.
In 2007, the complete nuclear genome sequence of "C. reinhardtii" was published.
Channelrhodopsin-1 and Channelrhodopsin-2, proteins that function as light-gated cation channels, were originally isolated from "C. reinhardtii". These proteins and others like them are increasingly widely used in the field of optogenetics.
Mitochondrial Significance.
The genome of C. Reinhardtii is significant for mitochondrial study as it is one species where the genes for 6 of the 13 proteins encoded for the mitochondria are found in the nucleus of the cell, leaving 7 in the mitochondria. In all other species these genes are present only in the mitochondria and are unable to be allotopically expressed. This is significant for the testing and development of therapies for genetic mitochondrial diseases.
Reproduction.
Vegetative cells of the "reinhardtii" species are haploid with 17 small chromosomes. Under nitrogen starvation, vegetative cells differentiate into haploid gametes. There are two mating types, identical in appearance, thus isogamous, and known as mt(+) and mt(-), which can fuse to form a diploid zygote. The zygote is not flagellated, and it serves as a dormant form of the species in the soil. In the light, the zygote undergoes meiosis and releases four flagellated haploid cells that resume the vegetative lifecycle.
Under ideal growth conditions, cells may sometimes undergo two or three rounds of mitosis before the daughter cells are released from the old cell wall into the medium. Thus, a single growth step may result in 4 or 8 daughter cells per mother cell.
The cell cycle of this unicellular green algae can be synchronized by alternating periods of light and dark. The growth phase is dependent on light, whereas, after a point designated as the transition or commitment point, processes are light-independent.
Eye spot.
Chlamydomonas reinhardtii has an eyespot similar to that of dinoflagellates(1). The eyespot is located near the cell equator. It is composed of a carotenoid-rich granule layer in the chloroplast which act like a light reflector.(2) The main function of the eyespot is the phototaxis, which consist of the movement (with the flagella) related to a light stimulus (3). The phototaxis is crucial for the alga and allows for localization of the environnement with optimal light conditions for photosynthesis. (4) Phototaxis can be positive or negative depending on the light intensity (1) . 
The phototactic pathway consists of four steps leading to a change in the beating balance between the two flagella (the cis-flagellum which is the one closest to the eyespot, and the trans-flagellum which is the one farthest from the eyespot). (5)
(1)Noriko Ueki, Takahiro Ide, Shota Mochiji, Yuki Kobayashi, Ryutaro Tokutsu, Norikazu Ohnishi, Katsushi Yamaguchi, Shuji Shigenobu, Kan Tanaka, Jun Minagawa, Toru Hisabori, Masafumi Hirono, Ken-ichi Wakabayashi, (2016) Eyespot-dependent determination of the phototactic sign in Chlamydomonas reinhardtii. , 113 (19) 5299-530
(2)Foster KW, Smyth RD (1980) Light Antennas in phototactic algae. Microbiol Rev 44(4): 572–630.
(3)Hegemann P, Berthold P (2009) Sensory photoreceptors and light control of flagellar activity.The Chlamydomonas Sourcebook, ed Witman G-B (Academic, Oxford), 2nd Ed, Vol 3, pp 395–430.
(4)Demmig-Adams B, Adams WW., III Photoprotection and other responses of plants to high light stress. Annu Rev Plant Physiol Plant Mol Biol. 1992;43:599–626.
(5)Hegemann P, Berthold P (2009) Sensory photoreceptors and light control of flagellar activity. The Chlamydomonas Sourcebook, ed Witman G-B (Academic, Oxford), 2nd, Ed, Vol 3, pp 395–430
Genetics.
The attractiveness of the algae as a model organism has recently increased with the release of several genomic resources to the public domain. The Chlre3 draft of the "Chlamydomonas" nuclear genome sequence prepared by Joint Genome Institute of the U.S. Dept of Energy comprises 1557 scaffolds totaling 120 Mb. Roughly half of the genome is contained in 24 scaffolds all at least 1.6 Mb in length. The current assembly of the nuclear genome is available online.
The ~15.8 Kb mitochondrial genome (database accession: NC_001638) is available online at the NCBI database. The complete ~203.8 Kb chloroplast genome (database accession: NC_005353) is available online.
In addition to genomic sequence data, there is a large supply of expression sequence data available as cDNA libraries and expressed sequence tags (ESTs). Seven cDNA libraries are available online. A BAC library can be purchased from the Clemson University Genomics Institute. There are also two databases of >50 000 and >160 000 ESTs available online.
A genome-wide collection of mutants with mapped insertion sites covering most nuclear genes is available: https://www.chlamylibrary.org/.
The genome of "C. reinhardtii" has been shown to contain N6-Methyldeoxyadenosine (6mA), a mark common in prokaryotes but much rarer in eukaryotes. Some research has indicated that 6mA in "Chlamydomonas" may be involved in nucleosome positioning, as it is present in the linker regions between nucleosomes as well as near the transcription start sites of actively transcribed genes.
Evolution.
"Chlamydomonas" has been used to study different aspects of evolutionary biology and ecology. It is an organism of choice for many selection experiments because (1) it has a short generation time, (2) it is both a autotroph and a facultative [[heterotroph], (3) it can reproduce both sexually and asexually, and (4) there is a wealth of genetic information already available.
Some examples (nonexhaustive) of evolutionary work done with "Chlamydomonas" include the evolution of sexual reproduction, the fitness effect of mutations, and the effect of adaptation to different levels of CO2.
According to one frequently cited theoretical hypothesis, sexual reproduction (in contrast to asexual reproduction) is adaptively maintained in benign environments because it reduces mutational load by combining deleterious mutations from different lines of descent and increases mean fitness. However, in a long-term experimental study of "C. reinhardtii", evidence was obtained that contradicted this hypothesis. In sexual populations, mutation clearance was not found to occur and fitness was not found to increase.
Motion.
[[File:Chlamydomonas_reinhardtii_trajectory.png|thumb]]Chlamydomonas Reinhardtii swims thanks to its two flagella, in a movement analogous to human [[breaststroke]]. Repeating this elementary movement 50 times per second the algae have a mean velocity of 70 µm/s; the genetic diversity of the different strains results in a huge range of values for this quantity. After few seconds of run, an asynchronous beating of the two flagella leads to a random change of direction. This movement is baptized "Run and Tumble". At a larger time and space scale, the random mouvement of the alga can be described as an active [[diffusion]] phenomenon.
DNA transformation techniques.
Gene transformation occurs mainly by homologous recombination in the chloroplast and heterologous recombination in the nucleus. The "C. reinhardtii" chloroplast genome can be transformed using microprojectile particle bombardment or glass bead agitation, however this last method is far less efficient. The nuclear genome has been transformed with both glass bead agitation and electroporation. The biolistic procedure appears to be the most efficient way of introducing DNA into the chloroplast genome. This is probably because the chloroplast occupies over half of the volume of the cell providing the microprojectile with a large target. Electroporation has been shown to be the most efficient way of introducing DNA into the nuclear genome with maximum transformation frequencies two orders of magnitude higher than obtained using glass bead method.
Production of biopharmaceuticals.
Genetically engineered "Chlamydomonas reinhardtii" has been used to produce a mammalian serum amyloid protein, a human [[antibody]] protein, human [[Vascular endothelial growth factor]], a potential [[HPV vaccine#Therapeutic HPV vaccines|therapeutic Human Papillomavirus 16 vaccine]], a potential [[malaria vaccine]] (an [[edible algae vaccine]]), and a complex designer drug that could be used to treat cancer.
Clean source of hydrogen production.
In 1939, the German researcher [[Hans Gaffron]] (1902–1979), who was at that time attached to the University of Chicago, discovered the hydrogen metabolism of unicellular green algae. "Chlamydomonas reinhardtii" and some other green algae can, under specified circumstances, stop producing oxygen and convert instead to the production of hydrogen. This reaction by [[hydrogenase]], an [[enzyme]] active only in the absence of oxygen, is short-lived. Over the next thirty years, Gaffron and his team worked out the basic mechanics of this photosynthetic hydrogen production by algae.
To increase the production of hydrogen, several tracks are being followed by the researchers.
Further reading.
Aoyama, H., Kuroiwa, T. and Nakamura, S. 2009. The dynamic behaviour of mitochondria in living zygotes during maturation and meiosis in "Chlamydomonas reinhardtii". "Eur. J. Phycol." 44: 497 - 507.
Jamers, A., Lenjou, M., Deraedt, P., van Bockstaele, D., Blust, R. and de Coen, W. 2009. Flow cytometric analysis of the cadmium-exposed green algae "Chlamydomonas reinhadtii" (Chlorophyceae). "Eur. J. Phycol." 44: 54 - 550.
External links.
[[Category:Chlamydomonadales]]
[[Category:Plant models]]
[[Category:Hydrogen production]]