A., Flexas, J. The results here indicate that the formation of a condensate is primarily due to the expression of the mature variant of EPYC1 rather than the level of expression per se, and that the required stoichiometry with Rubisco is achievable in planta. Bot. A Do-It-Yourself phenotyping system: measuring growth and morphology throughout the diel cycle in rosette shaped plants. These cookies will be stored in your browser only with your consent. Rosette growth rates were quantified using an in-house imaging system45. Roles PLOS ONE promises fair, rigorous peer review, generating a pyrenoid-like condensate in a plant chloroplast provides a platform for introducing bicarbonate (hco 3) channels/pumps at the chloroplast envelope (e.g. Sci. Sci. https://doi.org/10.1371/journal.pone.0185039.s005. 214, 655667 (2017). Rubisco small-subunit alpha-helices control pyrenoid formation in Chlamydomonas. 37, 19892001 (2014). What is the formula for calculating solute potential? This consisted of a binary vector containing two gene expression cassettes, each encoding mature EPYC1 with an Arabidopsis chloroplastic signal peptide and fused to a different version of GFP (turboGFP (tGFP) or enhanced GFP (eGFP)) to avoid possible recombination events (Fig. Cite this article. There is only one subunit of PSII, and no subunits of the two remaining major complexes; the cytochrome b6/f complex, and ATP synthase (Fig 3, S3 Table). Each cell contains a distinct nucleus, a central vacuole, and a large thin chloroplast with at least one pyrenoid. et al. The immunoblots shown were derived from the same experiment and gels/blots were processed in parallel. and A.J.M. Salesse-Smith, C. E. et al. In: Ehleringer J, Hall A, Farquhar G, (eds) Stable isotopes and plant carbon-water relations. Yamano, T., Sato, E., Iguchi, H., Fukuda, Y. . In addition, our quantitative proteomic analysis revealed that 85 proteins were strongly enriched in P fractions from WT compared to P fractions from rbcL and SSAT strains. Writing review & editing, Roles https://doi.org/10.1126/sciadv.abd2408 (2020). Anyone you share the following link with will be able to read this content: Sorry, a shareable link is not currently available for this article. This work used infrastructure in the Center for Microscopy and Cell Imaging and the Center for Structural and Functional Genomics, both at Concordia University. The dense cytoplasm is filled up in the cell. Yamano, T. et al. Five proteins function in starch degradation: an alpha-amylase, two isoamylases (ISA1-2) and two alpha-glucan water dikinases (GWD1-2) [12]. Methodology, The pyrenoid is a microcompartment within the chloroplasts of algae and hornworts. Level 2 vectors were transformed into Agrobacterium tumefaciens (AGL1) for stable insertion in Arabidopsis plants by floral dipping40. Overexpression of Rubisco subunits with RAF1 increases Rubisco content in maize. Moreover, proteins of the pyrenoid-deficient control strains that are localized to the pyrenoid in WT cells could have incidentally fractionated to the control P fractions by virtue of their physicochemical properties or presence in supramolecular assemblies and therefore may have been subtracted inappropriately from the proteome. Since fraction I protein is large, it seems reasonable to assume that this protein provides the granular matrix of the pyrenoid between the pyrenoid thylakoids. Similarly, our pyrenoid proteome does not resemble the proteome of the chloroplast stroma in C. reinhardtii; it contains only 14 proteins of the 274 reported in the stroma proteome (S3 Fig) [2]. a EPYC1 and Rubisco protein levels in whole leaf tissue (input), the supernatant following condensate extraction and centrifugation (supernatant) and the pellet (pellet) as assessed by immunoblot analyses with anti-EPYC1, anti-Rubisco (LSU and SSU shown) or anti-CrRbcS2 antibodies. Plant Cell Environ. Pyrenoid protein bodies are associated with starch storage in algae and hornworts. Together, these results suggest that carboxysomes and pyrenoids have analogous functions relating to gene expression, in addition to their known common role in CO2 fixation. Detergent-solubilized material remained in the supernatant (S). J. Exp. Immunoblots results were representative of four gels/blots for EPYC1, two for Rubisco, and one for CrRbcS2. volume11, Articlenumber:6303 (2020) Two papers from the Jonikas lab in this issue of Cell provide new insights into the structure, protein composition, and dynamics of this important organelle. Samples are shown for WT plants (WT), and S2Cr mutants not expressing EPYC1 (S2Cr) and expressing EPYC1-dGFP (S2Cr_EPYC1). The pyrenoid is a membrane-less organelle that exists in various photosynthetic organisms, such as algae, and wherein most global CO2 fixation occurs. Proc. Software, Biochemical and proteomic analyses of pyrenoids have been hampered by their instability during cell breakage and fractionation [10]. Writing review & editing, Affiliation https://doi.org/10.1007/BF00208310. lcia, shown in red) 34. Acs Synth. FRAP was carried out on live leaf tissue and tissue that had been fixed by infiltrating with 4% (v/v) formaldehyde for 90min in a vacuum chamber, then infiltrating three times for 5min each with phosphate-buffered saline (PBS) at pH 7.4. Plant Biotechnol. Using radioisotope pulse labeling experiments, we show that pyrenoid-associated ribosomes could be engaged in the localized synthesis of the large subunit of Rubisco. 4, 5). The datasets and plant materials generated and analysed during the current study are available from the corresponding author upon request. Gas exchange data were corrected for CO2 diffusion from the measuring chamber as in Bellasio et al.48. (C and D) Results of SDS-PAGE and silver-staining reveal proteins of the P and S fractions from (C) cells and isolated chloroplasts and (D) WT and the pyrenoid-deficient control strains SSAT and rbcL. Pyrenoids are sub-cellular micro-compartments found in chloroplasts of many algae, and in a single group of land plants, the hornworts. Accurate normalization of real-time quantitative RT-PCR data by geometric averaging of multiple internal control genes. This pyrenoid suspension was incubated with Alexa Fluor 488 AffiniPure Goat Anti-Rabbit IgG secondary antibody (1:200, Invitrogen) for 45 min at room temperature with gentle agitation. Gene-specific primers were designed to amplify either eGFP (5-CAGATTACGCCGTGAGCAAG-3 and 5-GCTGAACTTGTGGCCGTTTA-3) or tGFP (5-GCATCAGGGGTCTTGAAAGC-3 and 5-TCCTTCAAAACGGTGGACCT-3) (IDT, Belgium). Source data are provided as a Source Data file. BST13, shown in orange)35, a lumenal carbonic anhydrase to convert HCO3 to CO2 for release into the surrounding Rubisco condensate (CAH3, shown in blue)36, mechanisms to capture CO2 as HCO3 (LCIB and LCIC, shown in purple)37,38 and traversing thylakoid membranes33. Nevertheless, our P fractions contained non-pyrenoid material, seen as structures that did not IF-stain for RbcL or RbcS (Fig 2). 17, 12081216 (1998). 51, 14531468 (2010). Fiji: an open-source platform for biological-image analysis. SIM image processing, reconstruction and analysis were carried out using the N-SIM module of the NIS-Element Advanced Research software. here. Two caveats should be considered. The intensity of the GFP fluorescence signal over the cross-section of a chloroplast is shown in WT (n=28), S2Cr (line Ep3, n=17) and 1AAtMOD (n=22) backgrounds. Proteins were validated if they were identified with at least two different peptides passing the peptide FDR filter. For the latter expression cassette, a dual terminator system was used to increase expression53 (Supplementary Fig. No, Is the Subject Area "Proteomics" applicable to this article? McGrath, J. M. & Long, S. P. Can the cyanobacterial carbon-concentrating mechanism increase photosynthesis in crop species? Long, S. P., Marshall-Colon, A. The number of copies varies between species; however, the pea chloroplasts from mature leaves normally contain about 14 copies of the genome. For each experiment, n 20 cells. Atkinson, N. et al. Its main function is it is the centre of carbon dioxide fixation. https://doi.org/10.1371/journal.pone.0185039.s006. However, expression of the full coding sequence of EPYC1 in Arabidopsis with plant-algal hybrid Rubisco did not result in Rubisco condensation18. The response of A to the intercellular CO2 concentration (Ci) was measured at various CO2 concentrations (50, 100, 150, 200, 250, 300, 350, 400, 600, 800, 1000 and 1200molmol1) under saturating light (1500molphotonsm2s1). Plants were cultivated under light levels typical for Arabidopsis growth (200molphotonsm2s1) or under higher light levels (900molphotonsm2s1) where growth rates are more limited by Rubisco activity (Figs. Read More. -3 Current address: Department of Molecular and Cellular Biology, Science Complex University of Guelph, Guelph, Ontario, Canada, Roles Publishers note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. These observations have significant evolutionary implications for the development of terrestrial photosynthesis during the colonisation of the land, raising the intriguing question of why the pyrenoid-based CO2-concentrating mechanism did not persist in the terrestrial environment. Investigation, Source data are provided as a Source Data file. Carousel with three slides shown at a time. No, Is the Subject Area "Carbon dioxide" applicable to this article? The mass spectrometry proteomics data are summarized in S1 Table and have been deposited to the ProteomeXchange Consortium via the PRIDE partner repository with the dataset identifier PXD004509 (http://proteomecentral.proteomexchange.org; Reviewer account details: Username: reviewer15271@ebi.ac.uk; Password: 9fcT5WTO). He, S. et al. Pyrenoid pellets were washed with prechilled acetone and maintained at -20C for at least 2 hours. Each cell has generally cup-shaped chloroplast, one eye-spot and two contractile vacuoles. Alcalde, M.) Ch. No, Is the Subject Area "Starches" applicable to this article? The occurrence of the chloroplast pyrenoid is correlated with the activity of a CO 2 -concentrating mechanism and carbon isotope discrimination in lichens and bryophytes The organic-matter carbon isotope discrimination () of lichens with a wide range of photobiont and/or cyanobiont associations was used to determine t. Condensate and chloroplast dimensions were measured from confocal images using Fiji (ImageJ, v1.52n)51. Therefore, it was necessary to use additional criteria to specifically identify pyrenoid proteins in P fractions. This suggests that the modest reductions in Rubisco turnover rate (kcatc) and specificity (SC/O) for the plant-algal hybrid Rubisco in S2Cr compared to WT plants have only a mild impact on the efficiency of photosynthetic CO2 assimilation and that the observed differences in growth rates are more associated with the reduced levels of Rubisco in S2Cr plants21. Meyer, M. T. et al. 115, 153166 (2013). We next tested if the condensates exhibited internal mixing characteristics consistent with the liquid-like behaviour of pyrenoids (Fig. The supernatant (S) and pellet (P) fractions can be stored at -80C. Natl Acad. Genome Biol. In other algae, pyrenoids are the sites of carbohydrate (typically starch) storage. Pyrenoid-enriched fractions were subjected to a quantitative proteomic analysis using accurate mass/high resolution tandem mass spectrometry (ESI-MS/MS). This work was supported by the Natural Environment Research Council (GR3/8813) and the Leverhulme Trust. The resulting transgenic plants (Ep) expressed both EPYC1::eGFP and EPYC1::tGFP, of which the latter was generally more abundant at the protein level (Fig. Data curation, Cell 171, 133147.e114 (2017). Analysis by SDS-PAGE and silver staining revealed that P fractions from rbcL lacked both RbcL and RbcS, which is known to have a very low abundance in RbcL-deficient mutants [25] (Fig 1D). First, these ribosomes could be bound to the chloroplast translation membranes that were detected adjacent to the pyrenoid, possibly physically connected to it, thereby resulting in their co-purification with pyrenoids. This probably corresponds to newly synthesized RbcL that pelleted due to close association with pyrenoids. Planta 190, 332345 (1993). Which is part of the pyrenoid matrix is traversed? Google Scholar. Large white structures are starch granules. EPYC1 was further fused at the C-terminus to either enhanced GFP (eGFP) or turboGFP (tGFP), and driven by the 35S CaMV promoter (35S prom) or CsVMV promoter (CsVMV prom), respectively. Letters indicate significant difference (p<0.05) of EpWT lines compared to Ep lines as determined by one-way ANOVA followed by Tukeys honestly significant difference (HSD) post-hoc tests (a, d). Martinus Nijhof, Dordrecht, pp 601609, Badger MR, Pfanz H, Buedel B, Heber U, Lange OL (1993) Evidence for the functioning of photosynthetic carbon dioxide concentrating mechanisms in lichens containing green algal and cyanobacterial photobionts. 3c)17. Similarly, T2 EPYC1-dGFP WT plants (EpWT) showed no significant differences compared to T2 segregant lines (AzWT). 158, 905916 (2012). Finally, results of MS analysis of the P fractions from isolated chloroplasts revealed primarily proteins that are known or predicted to be in the chloroplast (S1 Fig). The present study set out to determine the cause of photosynthetic limitation in these pyrenoid-less lines. Subsequent Coomassie staining of denatured, gel-separated extracts revealed that nearly half (49%) of Rubisco in the initial extract contained Chlamydomonas SSU, while 82% of Rubisco in the pelleted condensate contained Chlamydomonas SSU (Fig. New pyrenoid functions are supported by proteins in tetrapyrrole and chlorophyll synthesis, carotenoid metabolism or amino acid metabolism. The major difference between floridean starch and starch is that floridean starch is generated in the cytoplasm and starch in plastids [44,52]. Nature Communications (Nat Commun) Processing of images was done with Leica LAS AF Lite software. a Fluorescence distribution plots of EPYC1-dGFP across the chloroplast. Dobrescu, A., Scorza, L. C. T., Tsaftaris, S. A. Pyrenoids therefore seem to have a role analogous to that of carboxysomes in cyanobacteria. What is a Pyrenoid and what does it do? USA 79, 69036907 (1982). Visualization, Traffic 20, 380389 (2019). 3. e Chlorophyll autofluorescence is reduced at the site of EPYC1-dGFP accumulation (white arrow). New Phytol. The cookie is used to store the user consent for the cookies in the category "Other. J. Exp. Formal analysis, From these chloroplasts, P fractions were prepared and subjected to analysis by SDS-PAGE and silver-staining (Fig 1C). a Schematic representation of the dual GFP expression system (EPYC1-dGFP) for EPYC1 truncated at amino acid residue 27 (as indicated by yellow triangles) and fused at the N-terminus to the chloroplastic transit peptide (TP) sequence of the Arabidopsis Rubisco small subunit RbcS1A. You are using a browser version with limited support for CSS. . Robert Spreitzer and Genhai Zhu, University of Nebraska) which was crossed into a CW15 genetic background in order to weaken its cell wall for lysis in Triton X-100, as part of the pyreniod enrichment protocol described in the next subsection. Finally, the pellet was resuspended in 25l of extraction buffer and used in confocal analysis or SDS-PAGE electrophoresis. Cells were not deprived of SO4 for more than the ~15 min required for centrifugation and resuspension and treatment with cycloheximide prior to pulse-labeling. This is your one-stop encyclopedia that has numerous frequently asked questions answered. . Proteins involved in CO2 fixation, CO2 metabolism (such as carbonic anhydrases) or annotated as `low CO2 inducible`genes are included in node Photosynthesis and CO2 metabolism. For comparisons of different genotypes, plants were grown from seeds of the same age and storage history, harvested from plants grown in the same environmental conditions. PubMedGoogle Scholar. Carboxysome encapsulation of the CO2-fixing enzyme Rubisco in tobacco chloroplasts. J Hattori Bot Lab 64: 7186, Kuchitsu K, Tsuzuki M, Miyachi S (1991) Polypeptide composition and enzyme activities of the pyrenoid and its regulation by CO2 concentrations in unicellular green algae. Software, 35S-pulse-labeling was performed for 5 min with 1.2 x 107 cells in 0.3 ml of TAP medium lacking NH4SO4. The chloroplast pyrenoid, an important component of the CO2 concentrating mechanism of algae, is a structure composed primarily of Rubisco. This plastids called chloroplast. Google Scholar, Green TGA, Lange OL (1994) Photosynthesis in poikilohydric plants: A comparison of lichens and bryophytes. Such a XIC-based relative quantification strategy allowed us to compare abundance values of proteins present in the different P fractions and determine proteins that are specific to the pyrenoid from those that can be considered as contaminants. type of starch molecule (Floridean starch) that is more highly branched than amylopectin. Shown are three T2 S2Cr transgenic plants expressing EPYC1-dGFP (Ep13) and azygous segregants (Az13), and S2Cr plants transformed with only EPYC1::tGFP (55.4kDa) or EPYC1::eGFP (63.9kDa). Comparative measurements of pyrenoid area were performed using Fiji on TEM cross-section images of WT C. reinhardtii cells (cMJ030) as described in Itakura et al.23. In protoplasts of leaf mesophyll cells, a condensate was visible in each chloroplast (Supplementary Fig. For all gas exchange experiments, the flow rate was kept at 200molmol1, leaf temperature was controlled at 25C and chamber relative humidity was maintained at ca. For example, two of the most abundant proteins of the pyrenoid, RbcL and Rubisco activase (RCA1) belong to this list of 85 proteins and would have been missed without quantitative analysis. J. Condensate formation in the 1AAtMOD background, where catalytic characteristics of the hybrid Rubisco are indistinguishable from that of WT Rubisco, indicates that the SSU can be further engineered to optimise phase separation, and Rubisco content and performance (Fig. Writing review & editing. D 74, 19 (2018). In summary, we have improved existing protocols for the purification of chloroplasts and pyrenoids. Nat. Nat. Cell 161, 5666 (2015). Its main function is it is the centre of carbon dioxide fixation. Natl Acad. In contrast to the present study, they did not report a functional characterization of the pyrenoid but focused on the characterization of a major pyrenoid protein, EPYC1, revealing it has a structural role in linking Rubisco holoenzyme complexes within the pyrenoid [18]. Shimada, T. L., Shimada, T. & Hara-Nishimura, I. In addition, the surface of the pyrenoid could serve as a platform for translation because the rbcL mRNA was seen to localize at the pyrenoid periphery in a translation-dependent manner [39] and we detected a newly synthesized protein, most probably RbcL, in P fractions, supporting the pyrenoid as a location of RbcL synthesis (Fig 4). The cookies is used to store the user consent for the cookies in the category "Necessary". However, a few concerns remained. Planta 239, 357366 (2014). The pyrenoidal linker protein EPYC1 phase separates with hybrid Arabidopsis-Chlamydomonas Rubisco through interactions with the algal Rubisco small subunit. 1 m that IF stain for marker proteins for the pyrenoid; RbcL and RbcS. Starch, a storage form of glucose, is often found around pyrenoids. . Correspondence to Biogenic and metabolic processes are localized to specialized domains or microcompartments to enhance their efficiency and suppress deleterious side reactions. In addition, the pyrenoid is believed to sequester Rubisco from O2 produced by photosystem II (PSII) complexes in thylakoid membranes located throughout the rest of the chloroplast [10].The pyrenoid is functionally analogous to the carboxysomes of cyanobacteria, in that both microcompartments provide Rubisco with a low O2/CO2 ratio to favor its carboxylase activity [11]. The condensates were spherical in shape and displaced native chlorophyll autofluorescence (Fig. Proteomics grade endoproteinases (Lys-C and Trypsin Gold) and ProteaseMax surfactant were purchased from Promega (Charbonnires, France). The pellet was washed twice with BB, resuspended in 100 l BB, and then immuno-reacted with affinity-purified antibodies against RbcL or RbcS (1:1,000 dilutions, Dr. Spreitzer, University of Nebraska) for 75 min at room temperature with gentle agitation. The Rubisco deficiency of rbcL considerably alters cellular metabolism and might therefore alter the non-pyrenoid proteins in the P fraction, thereby biasing the subtraction of contaminants. In: Schulze E, Caldwell MD (eds) Ecophysiology of photosynthesis. Differential tissue-specific expression of NtAQP1 in Arabidopsis thaliana reveals a role for this protein in stomatal and mesophyll conductance of CO2 under standard and salt-stress conditions. Decreased ribulose-1,5-bisphosphate carboxylase-oxygenase in transgenic tobacco transformed with antisense rbcS. The 488nm laser was set to 2% power for pre- and post-bleach images, and 25% for the bleaching step. 9 (CSIRO Publishing, 2019). To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/. Nevertheless, more than one third of the proteins are predicted to be localized outside of chloroplasts even though the samples were prepared from purified chloroplasts. A permeable sub-cellular micro-compartment surrounded by the layers of starch that facilitates photosynthetic CO2 fixation by ribulose-1,5-bisphosphate carboxylase (RuBisCO) in the chloroplast of certain algae and land plants are called pyrenoid protein bodies. Pyrenoids are associated with the operation of a carbon-concentrating mechanism (CCM). During normal plastid development in Euglena, the pyrenoid differentiates between 24 and 48 hours of illumination (Klein et al, 1972; Ben-Shaul et al, 1964) (Fig. Also displayed are a T2 EPYC1-dGFP WT transformant (EpWT) and azygous segregant (AzWT). b Coomassie-stained SDS-PAGE gel showing the composition of the pelleted condensate. 2A-K ), along with cell size and apical shapes ( Figs. non-truncated) variant of EPYC1-dGFP in Arabidopsis chloroplasts previously did not result in phase separation18, which was attributed to low levels of expression and an incompatible stoichiometry between EPYC1 and Rubisco, and possible proteolytic degradation. Methodology, Price, G. D., Badger, M. R. & von Caemmerer, S. The prospect of using cyanobacterial bicarbonate transporters to improve leaf photosynthesis in C3 crop plants. The condensates could be pelleted by gentle centrifugation16, and isolated condensates were shown to be enriched in EPYC1-dGFP and Rubisco (Fig. The numerous small spherical blobs along with the corners of the chloroplast are known as the pyrenoids. Moreover, previous results based on histochemical staining and electron microscopy suggest pyrenoid tubules in red algal species lack PSII [10]. & Sanchez-Baracaldo, P. The possible evolution and future of CO2-concentrating mechanisms. Dang., there is a marked difference in density between the chloroplast matrix and the pyrenoid, with an abrupt transition from the one to the other (Sager & Palade, 1957). Immunoblot analyses revealed that the isolated chloroplasts were, relative to total cellular protein content, depleted at least 7-fold of endoplasmic reticulum (BIP), 15-fold of mitochondria (AOX1), and 500-fold of cytoplasm (CyL4) (Fig 1B). In addition, nine proteins have predicted starch-related functions based on their sequences which contain an alpha amylase catalytic domain or a starch-binding domain. Formal analysis, 61, 519528 (2010). Conceptualization, 51, 659668 (2000). 1c and2a). Software, Validation, While most algae have 1 pyrenoid per chloroplast, here we describe a mutant in the model alga Chlamydomonas that has on average 10 pyrenoids per chloroplast. Pyrenoids contain protein and starch. This microcompartment enhances the photosynthetic CO2-fixing activity of the Calvin-Benson cycle enzyme Rubisco, suppresses an energetically wasteful oxygenase activity of Rubisco, and mitigates limiting CO2 availability in aquatic environments. During normal plastid development in Euglena, the pyrenoid differentiates between 24 . 35S-labelled proteins were revealed by phosphorimaging. The pyrenoid is a proteinaceous structure found in the chloroplast of most unicellular algae. Its known function is to promote photosynthetic CO2 fixation by the enzyme By submitting a comment you agree to abide by our Terms and Community Guidelines. Out of these, the cookies that are categorized as necessary are stored on your browser as they are essential for the working of basic functionalities of the website. This plastid contains a starch storing region called pyrenoids. Scale bar=0.5m for both images. . Biochem.-Us 58, 33653376 (2019). Samples were heated at 70C for 15min with 1 Bolt LDS sample buffer (ThermoFisher Scientific, UK) and 200mM dithiothreitol (DTT). Pyrenoids are subcellular microcompartments present in the chloroplast of hornworts. Reversed phase C18 spin columns, precolumns and analytical columns were all obtained from Thermo Scientific (Les Ulis, France). Formal analysis, These results demonstrate that the chloroplast ribosomes did not pellet with pyrenoids due to an association with chloroplast translation membranes and they support the pyrenoid being the location of rbcL mRNA translation [39]. Conceptualization, It is found around the pyrenoid and might be involved in the prevention of CO 2 leakage from the pyrenoid to the chloroplast stroma [24,100]. Protoplasma 156: 117129, Vaughn KC, Ligrone R, Owen HA, Hasegawa J, Campbell EO, Renzaglia KS, Monge-Najera J (1992) The anthocerote chloroplast: a review. CAS These pyrenoid-enriched fractions were shown to contain both subunits of Rubisco and a starch synthase. Investigation, This pyrenoid proteome contains 190 proteins, many of which function in processes that are known or proposed to occur in pyrenoids: e.g. In Chlamydomonas, Rubisco in the pyrenoid is held together by the linker protein EPYC1. Plants 4, 802810 (2018). A spatial interactome reveals the protein organization of the algal CO2-concentrating mechanism. For each technical replicate, protein abundance values were calculated by summing abundance values of all unique and razor peptides available using the Protein Quantifier node and, for each biological replicate, protein abundance values consisted of the average of two technical replicates. The 81 proteins of unknown function reveal candidates for new participants in these processes. Supervision, Scale bar=1m for both images. Oecologia 54: 275280, Hassel de Menendez GG (1988) A proposal for a new classification of the genera within the Anthocerotophyta. In these algae, pyrenoids probably function to fix carbon. The 35S cauliflower mosaic virus (CaMV) promoter and CsVMV (cassava vein mosaic virus) promoter were used to drive expression. Furthermore, the observed reduction in proteolytic degradation may be accounted for by sequestration of EPYC1 within a phase-separated compartment (Fig. Investigation, eLife 4, e04889 (2015). LCIA, shown in red)34 and thylakoid membrane (e.g. Sci. Assembly of the algal CO2-fixing organelle, the pyrenoid, is guided by a Rubisco-binding motif. J Electron Microsc 41: 424433, Palmqvist K, Samuelsson G, Badger, MR (1994) Photobiont related differences in carbon acquisition among green-algal lichens.
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