Chlamydomonas reinhardtii

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Chlamydomonas reinhardtii
Chlamydomonas6-1.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Clade: Viridiplantae
Division: Chlorophyta
Class: Chlorophyceae
Order: Chlamydomonadales
Family: Chlamydomonadaceae
Genus: Chlamydomonas
Species:
C. reinhardtii
Binomial name
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 apparatus that senses light.

Contents

Chlamydomonas species are widely distributed worldwide in soil and fresh water, of which Chlamydomonas reinhardtii is one of the most common and widespread. [1] C. 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 collected near Amherst, Massachusetts, in 1945 by Gilbert M. Smith. [2] [3]

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. [4]

Description

Cells of Chlamydomonas reinhardtii are mostly spherical, but can range from ellipsoidal, ovoid, obovoid, or asymmetrical. They are 10–22 μm long and 8–22 μm wide. The cell wall is thin, lacking a papilla. The flagella are 1.5 to 2 times the length of the cell body. Cells contain a single cup-shaped chloroplast lining the bottom of the cell, with a single basal pyrenoid. [1]

Eye spot

C. reinhardtii has an eyespot apparatus similar to that of dinoflagellates. [5] 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. [6] The main function of the eyespot is the phototaxis, which consist of the movement (with the flagella) related to a light stimulus. [7] The phototaxis is crucial for the alga and allows for localization of the environment with optimal light conditions for photosynthesis. [8] Phototaxis can be positive or negative depending on the light intensity. [5] 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). [7]

Model organism

Cross section of a Chlamydomonas reinhardtii cell Cross section of a Chlamydomonas reinhardtii algae cell, a 3D representation.jpg
Cross section of a Chlamydomonas reinhardtii cell

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. [9]

Channelrhodopsin-1 and Channelrhodopsin-2, proteins that function as light-gated cation channels, were originally isolated from C. reinhardtii. [10] [11] These proteins and others like them are increasingly widely used in the field of optogenetics. [12]

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. [13] In species outside of Chlorophyceae, 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 reinhardtii species are haploid with 17 small chromosomes. Under nitrogen starvation, vegetative cells differentiate into haploid gametes. [14] 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. [15]

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. [16]

The ~15.8 Kb mitochondrial genome (database accession: NC_001638) is available online at the NCBI database. [17] The complete ~203.8 Kb chloroplast genome (database accession: NC_005353) is available online. [18] [19]

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. [20] A BAC library can be purchased from the Clemson University Genomics Institute. [21] There are also two databases of >50 000 [22] and >160 000 [23] ESTs available online.

A genome-wide collection of mutants with mapped insertion sites covering most nuclear genes [24] [25] is available: https://www.chlamylibrary.org/.

The genome of C. reinhardtii has been shown to contain N6-Methyladenosine (m6A), a mark common in prokaryotes but much rarer in eukaryotes. [26] 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. [27]

C. reinhardtii appears to be capable of several DNA repair processes. [28] These include recombinational repair, strand break repair and excision repair. In particular, C. reinhardtii chloroplasts possess an efficient system for repairing DNA double-strand breaks. [29] In chloroplast DNA homologous recombinational repair is strongly stimulated by double-strand breaks. [29]

Experimental 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 an 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, [30] the fitness effect of mutations, [31] and the effect of adaptation to different levels of CO2. [32]

According to one frequently cited theoretical hypothesis, [33] 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. [34]

Motion

C. reinhardtii trajectory, in HSA (culture medium), under red light. Chlamydomonas reinhardtii trajectory.png
C. reinhardtii trajectory, in HSA (culture medium), under red light.

C. reinhardtii swims thanks to its two flagella, [35] 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; [36] 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, a movement called "run and tumble". [35] At a larger time and space scale, the random movement of the alga can be described as an active diffusion phenomenon. [37]

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.[ citation needed ]

Practical uses

Production of biopharmaceuticals

Genetically engineered C. reinhardtii has been used to produce a mammalian serum amyloid protein (needs citation), a human antibody protein (needs citation), human Vascular endothelial growth factor, a potential therapeutic Human Papillomavirus 16 vaccine, [38] a potential malaria vaccine (an edible algae vaccine), [39] and a complex designer drug that could be used to treat cancer. [40]

Alternative protein source

C. reinhardtii has been suggested as a new algae-based nutritional source. Compared to Chlorella and Spirulina, C. reinhardtii was found to have more Alpha-linolenic acid, and a lower quantity of heavy metals while also containing all the essential amino acids and similar protein content. [41] Triton Algae Innovations was developing a commercial alternative protein product made from C reinhardtii.

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. C 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. [42]

To increase the production of hydrogen, several tracks are being followed by the researchers.

1. Photosystem II-dependent water oxidation reaction:    2H2O → 4H+ + O2 + 4e⁻;
2. Hydrogenase-dependent reversible reduction of protons to molecular hydrogen:    4H+ + 4e⁻ ⇄ 2H2.

If the water oxidation reaction leading to O2 production is balanced with O2 consumption, either by respiration or by introducing additional O2 absorbents or scavengers, the photosynthetic production of H2 can be sustained for a prolonged period. This balance prevents the inhibition of hydrogenase activity by accumulated O2, ensuring steady hydrogen production under these optimized conditions [50] .

See also

Related Research Articles

<span class="mw-page-title-main">Chloroplast</span> Plant organelle that conducts photosynthesis

A chloroplast is a type of organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. Chloroplasts have a high concentration of chlorophyll pigments which capture the energy from sunlight and convert it to chemical energy and release oxygen. The chemical energy created is then used to make sugar and other organic molecules from carbon dioxide in a process called the Calvin cycle. Chloroplasts carry out a number of other functions, including fatty acid synthesis, amino acid synthesis, and the immune response in plants. The number of chloroplasts per cell varies from one, in some unicellular algae, up to 100 in plants like Arabidopsis and wheat.

<span class="mw-page-title-main">Chlorophyta</span> Phylum of green algae

Chlorophyta is a division of green algae informally called chlorophytes.

<span class="mw-page-title-main">Symbiogenesis</span> Evolutionary theory holding that eukaryotic organelles evolved through symbiosis with prokaryotes

Symbiogenesis is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibly other organelles of eukaryotic cells are descended from formerly free-living prokaryotes taken one inside the other in endosymbiosis. Mitochondria appear to be phylogenetically related to Rickettsiales bacteria, while chloroplasts are thought to be related to cyanobacteria.

<span class="mw-page-title-main">Plastid</span> Plant cell organelles that perform photosynthesis and store starch

A plastid is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. Plastids are considered to be intracellular endosymbiotic cyanobacteria.

<i>Chlamydomonas</i> Genus of algae

Chlamydomonas is a genus of green algae consisting of about 150 species of unicellular flagellates, found in stagnant water and on damp soil, in freshwater, seawater, and even in snow as "snow algae". Chlamydomonas is used as a model organism for molecular biology, especially studies of flagellar motility and chloroplast dynamics, biogenesis, and genetics. One of the many striking features of Chlamydomonas is that it contains ion channels (channelrhodopsins) that are directly activated by light. Some regulatory systems of Chlamydomonas are more complex than their homologs in Gymnosperms, with evolutionarily related regulatory proteins being larger and containing additional domains.

<span class="mw-page-title-main">Pyrenoid</span> Organelle found within the chloroplasts of algae and hornworts

Pyrenoids are sub-cellular phase-separated micro-compartments found in chloroplasts of many algae, and in a single group of land plants, the hornworts. Pyrenoids are associated with the operation of a carbon-concentrating mechanism (CCM). Their main function is to act as centres of carbon dioxide (CO2) fixation, by generating and maintaining a CO2 rich environment around the photosynthetic enzyme ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO). Pyrenoids therefore seem to have a role analogous to that of carboxysomes in cyanobacteria.

A hydrogenase is an enzyme that catalyses the reversible oxidation of molecular hydrogen (H2), as shown below:

<span class="mw-page-title-main">Biohydrogen</span> Hydrogen that is produced biologically

Biohydrogen is H2 that is produced biologically. Interest is high in this technology because H2 is a clean fuel and can be readily produced from certain kinds of biomass, including biological waste. Furthermore some photosynthetic microorganisms are capable to produce H2 directly from water splitting using light as energy source.

<span class="mw-page-title-main">I-CreI</span> Homing endonuclease

I-CreI is a homing endonuclease whose gene was first discovered in the chloroplast genome of Chlamydomonas reinhardtii, a species of unicellular green algae. It is named for the facts that: it resides in an Intron; it was isolated from Clamydomonas reinhardtii; it was the first (I) such gene isolated from C. reinhardtii. Its gene resides in a group I intron in the 23S ribosomal RNA gene of the C. reinhardtii chloroplast, and I-CreI is only expressed when its mRNA is spliced from the primary transcript of the 23S gene. I-CreI enzyme, which functions as a homodimer, recognizes a 22-nucleotide sequence of duplex DNA and cleaves one phosphodiester bond on each strand at specific positions. I-CreI is a member of the LAGLIDADG family of homing endonucleases, all of which have a conserved LAGLIDADG amino acid motif that contributes to their associative domains and active sites. When the I-CreI-containing intron encounters a 23S allele lacking the intron, I-CreI enzyme "homes" in on the "intron-minus" allele of 23S and effects its parent intron's insertion into the intron-minus allele. Introns with this behavior are called mobile introns. Because I-CreI provides for its own propagation while conferring no benefit on its host, it is an example of selfish DNA.

<span class="mw-page-title-main">Eyespot apparatus</span> Photoreceptive organelle

The eyespot apparatus is a photoreceptive organelle found in the flagellate or (motile) cells of green algae and other unicellular photosynthetic organisms such as euglenids. It allows the cells to sense light direction and intensity and respond to it, prompting the organism to either swim towards the light, or away from it. A related response occurs when cells are briefly exposed to high light intensity, causing the cell to stop, briefly swim backwards, then change swimming direction. Eyespot-mediated light perception helps the cells in finding an environment with optimal light conditions for photosynthesis. Eyespots are the simplest and most common "eyes" found in nature, composed of photoreceptors and areas of bright orange-red red pigment granules. Signals relayed from the eyespot photoreceptors result in alteration of the beating pattern of the flagella, generating a phototactic response.

<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.

<span class="mw-page-title-main">Ferredoxin hydrogenase</span> Class of enzymes

In enzymology, ferredoxin hydrogenase, also referred to as [Fe-Fe]hydrogenase, H2 oxidizing hydrogenase, H2 producing hydrogenase, bidirectional hydrogenase, hydrogenase (ferredoxin), hydrogenlyase, and uptake hydrogenase, is found in Clostridium pasteurianum, Clostridium acetobutylicum,Chlamydomonas reinhardtii, and other organisms. The systematic name of this enzyme is hydrogen:ferredoxin oxidoreductase

<span class="mw-page-title-main">Phototaxis</span> Directed movement of a motile cell or organism in response to light

Phototaxis is a kind of taxis, or locomotory movement, that occurs when a whole organism moves towards or away from a stimulus of light. This is advantageous for phototrophic organisms as they can orient themselves most efficiently to receive light for photosynthesis. Phototaxis is called positive if the movement is in the direction of increasing light intensity and negative if the direction is opposite.

<i>Guillardia</i> Genus of single-celled organisms

Guillardia is a genus of marine biflagellate cryptomonad algae with a plastid obtained through secondary endosymbiosis of a red alga.

<i>Volvox carteri</i> Species of alga

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<span class="mw-page-title-main">Sabeeha Merchant</span> Plant scientist

Sabeeha Sabanali Merchant is a professor of plant biology at the University of California, Berkeley. She studies the photosynthetic metabolism and metalloenzymes In 2010 Merchant led the team that sequenced the Chlamydomonas genome. She was elected a member of the National Academy of Sciences in 2012.

<span class="mw-page-title-main">Protist locomotion</span> Motion system of a type of eukaryotic organism

Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly unicellular and microscopic. Many unicellular protists, particularly protozoans, are motile and can generate movement using flagella, cilia or pseudopods. Cells which use flagella for movement are usually referred to as flagellates, cells which use cilia are usually referred to as ciliates, and cells which use pseudopods are usually referred to as amoeba or amoeboids. Other protists are not motile, and consequently have no built-in movement mechanism.

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.

<i>Chlamydomonas moewusii</i> Species of algae

Chlamydomonas moewusii is a species of unicellular green alga belonging to the genus Chlamydomonas. C. moewusii is typically a freshwater species and occupies a significant position as a model organism for various scientific studies due to its relatively simple cellular structure and ease of cultivation.

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