Therefore, the dissection of a process in C. However, more recent higher-resolution imaging resolved Rubisco distribution within the matrix as liquid-like rose Freeman Rosenzweig et al. The same cannot be said for most eukaryotic systems that have retained cilia; mutations in human ciliary proteins lead to diseases referred to as ciliopathies that include male infertility, kidney and eye malfunction, polydactyly, and brain disorders. In Situ Electron Cryotomography By necessity, researchers tend to focus on a handful of genes to uncover the mechanisms controlling their expression and the functions of their products, with the ultimate goal of placing the encoded proteins into the much larger context of a living cell.
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Why not change that to the push of a button on a phone from wherever you are? We also present resources for current and future scientists who may wish to expand their studies to the realm of microalgae.
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Algae represent a very large and diverse polyphyletic group of photosynthetic eukaryotes Blaby-Haas and Merchant, They occupy all possible ecological niches on the planet, and therefore constitute a potential reservoir of untapped functional capabilities for adaptation to the environment. They also offer a low-cost option for the large-scale production of high-value molecules, since algae only require water, salts, air, and light. Unicellular algae such as the ciliated green alga Chlamydomonas reinhardtii offer high signal-to-noise during experiments due to the ease of growth in controlled medium and environments temperature and light regimes and the homogenous nature of the cultures, and they grow much more rapidly than classic plant models.
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With its haploid genome, C. Other topics of research using C. Responses to excess light and the dissipation of light energy to prevent cellular damage are another research avenue that has benefitted enormously from the analysis of C. Additional research questions being addressed by the study of C. Inside its pages and three volumes, The Chlamydomonas Sourcebook encompasses major research topics, history, and methodology Harris, We will feature selected examples below where C.
Despite conservation of the underlying proteins for example, cyclins, cyclin-dependent kinases, and the anaphase-promoting complexbudding yeast is evolutionarily distant from algae and plants.
This separation of phase along the diurnal cycle provides a predictable temporal cascade that starts with cell growth fueled by photosynthesisfollowed by the commitment to divide, resorption of cilia, doubling of DNA and histone contents, mitosis, and the growth of new cilia. With the help of robotics and semiautomated imaging, it is possible to identify temperature-sensitive mutants affected in some aspect of cell cycle progression, and whole-genome sequencing can be used to pinpoint the causal mutation.
A clear challenge is to understand the role of each cell cycle gene, but looking at their expression windows over the diurnal cycle should help narrow down the options Zones et al.
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Chloroplast Biogenesis and Photosynthesis C. Mutants with defects in the light reactions can be specifically enriched by the addition of the bactericidal agent metronidazole. Also known as Flagyl, this compound is reduced to its toxic form by ferredoxin; only cells that cannot reduce ferredoxin survive in its presence Schmidt et al.
Through classic genetic screens, acetate-requiring ac photosynthetic mutants were isolated based on the requirement for acetate as a reduced carbon source hence the name acetate requiring or increased chlorophyll fluorescence, a consequence of disrupted electron transport Levine, a ; Bennoun and Levine, ; Bennoun et al. Chlorophyll fluorescence reflects the progression of electron flow through the electron transport chain: its blockage increases fluorescence, as light energy can no longer be used for photochemistry.
The same fluorescence assay provides a powerful means to determine allelism and genomic rescue when identifying the underlying causal loci. In land plants, mutants in the homologous genes also display high chlorophyll fluorescence but are generally seedling-lethal and must be maintained as a segregating stock Maiwald et al.
A Series of Fortunate Events: Introducing Chlamydomonas as a Reference Organism
Homozygous mutant plants can, however, survive on medium supplemented with sucrose, which is taken up by dedicated transporters missing in Chlamydomonas.
Early evidence for nucleus- and plastid-encoded components of the photosynthetic apparatus came from radiolabeling C. Pulse-chase labeling of cells with 35S in the presence or absence of a plastid translation inhibitor also revealed that the small subunit of Rubisco translated in the cytoplasm is rapidly degraded when the large subunit is missing Schmidt and Mishkind,mirroring observations collected on photosystem II PSII proteins in the absence of plastid-encoded D1 and D2 proteins Erickson et al.
The control of epistasy of synthesis CES model describes how the translation potential of individual components of a photosynthetic protein complex depends on the presence of the other constituents Choquet and Wollman, The genetic dissection of photosynthesis owes much to the collection of ac mutants from Levine, followed by new mutants from career-long screens from the Rochaix and Wollman laboratories.
Analysis of some mutants provided experimental validation for trans-splicing of the plastid-encoded psaA gene Kück et al. Coordination between nucleus- and plastid-encoded subunits of photosynthetic complexes and enzymes is critical for proper chloroplast function. Such RNA binding proteins exhibit sequence specificity for a given chloroplast RNA and are encoded by large gene families in land plants that have not yet expanded in algae Jalal et al.
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Photosynthesis and State Transitions Analyses of other mutants have led to the genetic dissection of state transitions. Although state transitions were first discovered in other organisms, the biochemical underpinnings of this process were elucidated in C.
State transitions require cytochrome Cyt b6f to sense the redox status of the plastoquinone pool, as a Cyt b6f mutant is locked in state 1 and primarily accumulates unphosphorylated LHCII proteins Wollman and Lemaire, Mutants lacking state transitions display identical chlorophyll fluorescence whether the plastoquinone pool is highly oxidized state 1 or reduced state 2.
In the stt7 mutant for state-transition deficientLHCII proteins are only found in their nonphosphorylated form. The stt7 mutant can still grow in the absence of acetate, albeit at a slower rate, indicating that state transitions contribute to fitness Fleischmann et al.
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The interaction of Cyt b6f with STT7 is promoted by reduced plastoquinone and results in the autophosphorylation and activation of the kinase. The STT7 homolog in Arabidopsis, STN7, also catalyzes LHCII phosphorylation in response to the redox status of plastoquinone, and the stn7 mutant shows growth defects under a fluctuating light regime, which more accurately mimics real-life growth conditions Bellafiore et al.
Therefore, the dissection of a process in C. This process may allow an increase in photosynthetic yields by better recycling excess light photon energy away from dissipation.
Photosynthesis and the CCM Oxygen and carbon dioxide compete for the active site of Rubisco, causing a decrease in photosynthetic efficiency. Rubisco evolved under low-oxygen conditions, but oxygenic photosynthesis altered the surrounding environment.
Biological innovations increased local relative CO2 concentrations to enhance photosynthetic performance.
These innovations include carboxysomes in cyanobacteria, C4 metabolism in land plants, and the CCM in C. A functional CCM complex starts with the uptake of bicarbonate from the environment, its delivery to the pyrenoid a site of concentrated Rubiscoand its conversion to CO2 by carbonic anhydrases. The putative membrane-bound transporters responsible for inorganic carbon uptake, the carbonic anhydrases, and the transcriptional network controlled by inorganic carbon availability were identified through comparative transcriptome analysis of cells grown under low and high CO2 concentrations and through genetic screens Jungnick et al.
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In addition, the biochemical identification of proteins in purified pyrenoid components was validated genetically Mackinder et al. Most CCM components were tentatively arranged along the bicarbonate chemical gradient created within the cell from the cell wall to the pyrenoid matrix based on their localization Mackinder et al.
Surprisingly, proteins can localize to distinct layers and puncta within the pyrenoid, in addition to the typical matrix localization of Rubisco and the Rubisco binding Essential Pyrenoid Component 1 EPYC1. The direct observation of the Rubisco small subunit, RBCS1, fused to YFP also resolved the organization of the pyrenoid matrix as a liquid whose contents can mix internally, unlike the more rigid cyanobacterial carboxysomes Freeman Rosenzweig et al.
The introduction of the Rubisco linker protein EPYC1 into plants or a version that can interact with plant Rubisco may be an important step toward the engineering of plants to form pyrenoids to enhance photosynthetic output.
Channelrhodopsins and Optogenetics For a unicellular organism, C.