Chlamydomonas reinhardtii is an ideal model organism for studying the cytoplasmic preassembly of ciliary dyneins. However, under normal liquid-culture conditions, Chlamydomonas preassembly-deficient mutants often show no cilia or a small number of ciliated cells (i.e., low ciliary ratio), which hinders further analysis of both ciliary dyneins and the phenotypes of these mutants. In this brief report, we present a modified culture method for one of the Chlamydomonas preassembly-deficient mutants, pf23. This method enables researchers to obtain enough pf23 cilia for small-scale biochemical, biophysical, structural and phenotypic analyses.
","acknowledgements":"We would like to thank Drs. Kazuo Inaba (University of Tsukuba) and Takashi Ishikawa (Paul Scherrer Institute), and Mr. Yoshio Inomata/Akinobu Toratani/Motoki Nishizaka (Osaka University) for their help and advice regarding the phenotypic analyses of the pf23 mutant. We would also like to thank Dr. Winfield S. Sale (Emory University) for his support and careful reading of the manuscript.
","authors":[{"affiliations":["Osaka University, Osaka, Ōsaka, Japan"],"departments":["Department of Biological Sciences, Graduate School of Science"],"credit":["conceptualization","fundingAcquisition","writing_originalDraft","investigation","methodology"],"email":"yamamoto.ryousuke.sci@osaka-u.ac.jp","firstName":"Ryosuke","lastName":"Yamamoto","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["Osaka University, Osaka, Ōsaka, Japan"],"departments":["Department of Biological Sciences, Graduate School of Science"],"credit":["conceptualization","supervision","fundingAcquisition","writing_reviewEditing"],"email":"takahide.kon@bio.sci.osaka-u.ac.jp","firstName":"Takahide","lastName":"Kon","submittingAuthor":false,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":null,"extendedData":[],"funding":"R.Y is a recipient of JSPS Grant-in-Aid for Scientific Research (C)(JP24K09454), and T.K was a recipient of JSPS Grant-in-Aid for Scientific Research (B)(JP17H03665).
","image":{"url":"https://portal.micropublication.org/uploads/d942a7bd3ed090e6f2c85d90ba272c19.png"},"imageCaption":"This is a schematic illustration of the ciliation method for pf23, the Chlamydomonas preassembly-deficient mutant. The panels [(A) - (F)] in this figure are explained in the \"Methods\" section of the main text. In our laboratory, we check the ciliary growth of the pf23 cells transferred to the SG medium every day (panel E). As early as 1 day after transferring the cells, we often observed that they had grown sufficient cilia to perform small-scale experiments. This figure was created using PowerPoint (Microsoft).
","imageTitle":"Culture method for the Chlamydomonas pf23 mutant with difficulty growing cilia
","methods":"","reagents":"MUTANT | GENOTYPE | DEFECTS | AVAILABLE FROM |
pf23-1 | pf23 | Cytoplasmic preassembly of several ciliary dyneins (Huang et al., 1979; Yamamoto et al., 2017) | Chlamydomonas resource center (https://www.chlamycollection.org/) |
Chlamydomonas is an excellent model organism (Marshall, 2024; Ostrowski et al., 2011; Vincensini et al., 2011) for studying the composition and function of motile cilia, organelles that play vital roles in various important biological processes in eukaryotes, such as fertility and development (Satir & Christensen, 2007). In this regard, this tiny alga serves as an outstanding organism for studying the pathogenesis of primary ciliary dyskinesia (PCD), a human disease caused by deficient ciliary motility (Chodhari et al., 2004; Zariwala et al., 2011). Motile cilia are powered by motor-protein complexes called \"ciliary dyneins\" (Kamiya & Yagi, 2014; King et al., 2023; Yamamoto et al., 2021). The ciliary dyneins are preassembled in the cytoplasm before being transported into cilia (in a sequential process called \"preassembly\")(Desai et al., 2017; Mitchell & Yamamoto, 2023), and defects in this process also cause PCD in humans [e.g. (Mitchison et al., 2012; Omran et al., 2008; Tarkar et al., 2013)]. Across species, many Chlamydomonas preassembly-deficient mutants have also been identified [e.g. (Dean & Mitchell, 2013; Desai et al., 2015; Huang et al., 1979; Kamiya, 1988; Yamamoto et al., 2010)]. However, researchers often face a major challenge when studying the mutants: under the standard liquid tris-acetate-phosphate (TAP) medium condition (Gorman & Levine, 1965; Harris, 1989), these preassembly mutants often grow no or very short cilia, which are insufficient for various experimental analyses. Through testing several culture conditions, we found that, under a specific condition, a classical preassembly mutant, pf23 (Huang et al., 1979), tends to grow cilia more efficiently than under the standard liquid TAP medium condition. In this brief report, we describe a simple culture method that results in efficient ciliary growth in the pf23 mutant and provides enough cilia for small-scale biochemical, biophysical, structural and phenotypic analyses. In our laboratory, we mainly use this method, or modified versions of it, to isolate cilia from pf23.
Methods:
The protocol for our new culture method (Figure 1) is described as follows:
[0] (Before the experiment/culture) Make ~ 200 - 500 mL of the liquid SG [Sager and Granick, or minimal (M)] medium [originally reported in (Sager & Granick, 1953), see also (Harris, 1989) and the Chlamydomonas resource center (https://www.chlamycollection.org/) for basic/conventional compositions] in a conical flask. Place a magnetic stirrer in the SG medium, then autoclave the SG medium and cool it to room temperature. There is no need to prepare bubbling equipment for aeration.
[1] Inoculate the target strain (in this case, pf23) onto the ~ 5 - 10 solid TAP plates (Gorman & Levine, 1965; Harris, 1989), each measuring ~ 90 mm, and ensure the Chlamydomonas cells are distributed evenly across the plates (Figure 1A). Incubate the cells at room temperature under appropriate light (e.g. under a desk lamp) for ~ 3 - 7 days to allow them to grow well and turn the plates green (Figure 1B).
[2] Using an inoculation loop, scrape all the cells (cultured for ~ 3 - 7 days) from all the solid TAP plates and transfer them into the ~ 200 - 500 mL of the liquid SG medium (prepared in [0])(Figure 1C). Since all the cells from the ~ 5 - 10 solid TAP plates are in the liquid SG medium, the medium should be green to dark green.
[3] Gently mix the Chlamydomonas culture in the SG medium (Figure 1D) at room temperature under appropriate light (e.g. under a desk lamp) with the magnetic stirrer (placed in the SG medium in [0]). There is no need for bubbling/aeration. The stirring should be moderately strong enough to disrupt the mother cell walls of palmelloid cells (Iwasa & Murakami, 1968; Khona et al., 2016; Wingfield & Lechtreck, 2018) without disrupting the cells themselves.
[4] Keep the Chlamydomonas culture mixed in the SG medium with the stirrer under light. The cells will often grow cilia 1 - 3 days after being transferred to the liquid SG medium. Using conventional light microscopy (e.g. Olympus BX50, at ~ ×200 magnification), check if the cells are ciliated every day after transferring them to the SG medium (Figure 1E). Once the cells are sufficiently ciliated (e.g. ciliary ratio ≧ ~ 70%), deciliate them (Figure 1F) and collect the cilia using the standard procedure (Craige et al., 2013; Witman, 1986).
Note that all the procedures should be aseptic, and contamination of the Chlamydomonas cultures by bacteria or fungi, especially during mixing in the SG medium, should be avoided. With this method, one of the Chlamydomonas preassembly-deficient mutants, pf23, tends to grow cilia efficiently. In the past, researchers observed that some preassembly-deficient mutants tend to grow cilia at a late culture stage in the nutrient-rich media (including the TAP medium)[e.g. (Huang et al., 1979; Yamamoto et al., 2010)], i.e., when nutrients in the nutrient-rich media decrease from the initial high amounts. We incorporated this observation into this method, establishing an easy way to grow pf23 cilia. Due to the availability of reagents/chemicals as well as historical reasons, compositions of the SG medium somewhat vary between laboratories studying Chlamydomonas, but we believe most of these media would work well for this method. In addition, in this method, other nutrient-poor Chlamydomonas media [e.g. nitrate-as-nitrogen-source (N) medium (Sager & Granick, 1953), see also the Chlamydomonas resource center (https://www.chlamycollection.org/)] could potentially be used as an alternative to the SG medium.
Conclusion:
We have long used this method, or modified versions of it [e.g. usage of \"Hutner’s trace elements\" (Hutner et al., 1950) instead of the trace elements], to isolate sufficient amounts of cilia from pf23, the classical Chlamydomonas preassembly-deficient mutant. This method is simple and convenient, can be performed using standard equipment, and is workable in most laboratories studying Chlamydomonas. This method could also be applicable to other Chlamydomonas ciliary and/or preassembly-deficient mutants that grow very short or few cilia under the standard TAP medium condition, such as pf13 (Huang et al., 1979; Omran et al., 2008) and pf22 (Huang et al., 1979; Mitchison et al., 2012).
","references":[{"reference":"Chodhari, R., Mitchison, H. M., & Meeks, M. (2004). Cilia, primary ciliary dyskinesia and molecular genetics. Paediatr Respir Rev, 5(1), 69-76.
","pubmedId":"15222957","doi":"10.1016/j.prrv.2003.09.005"},{"reference":"Craige, B., Brown, J. M., & Witman, G. B. (2013). Isolation of Chlamydomonas flagella. Curr Protoc Cell Biol, Chapter 3 (Unit 3.41), 1-9.
","pubmedId":"23728744","doi":"10.1002/0471143030.cb0341s59"},{"reference":"Dean, A. B., & Mitchell, D. R. (2013). Chlamydomonas ODA10 is a conserved axonemal protein that plays a unique role in outer dynein arm assembly. Mol Biol Cell, 24(23), 3689-3696.
","pubmedId":"24088566","doi":"10.1091/mbc.E13-06-0310"},{"reference":"Desai, P. B., Dean, A. B., & Mitchell, D. R. (2017). Chapter 4 - Cytoplasmic preassembly and trafficking of axonemal dyneins. In S. M. King (Ed.), Dyneins (The Biology of Dynein Motors)( 2 nd Edition) (pp. 140-161). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-809471-6.00004-8"},{"reference":"Desai, P. B., Freshour, J. R., & Mitchell, D. R. (2015). Chlamydomonas axonemal dynein assembly locus ODA8 encodes a conserved flagellar protein needed for cytoplasmic maturation of outer dynein arm complexes. Cytoskeleton (Hoboken), 72(1), 16-28.
","pubmedId":"25558044","doi":"10.1002/cm.21206"},{"reference":"Gorman, D. S., & Levine, R. P. (1965). Cytochrome f and plastocyanin: their sequence in the photosynthetic electron transport chain of Chlamydomonas reinhardi. Proc Natl Acad Sci U S A, 54(6), 1665-1669.
","pubmedId":"4379719","doi":"10.1073/pnas.54.6.1665"},{"reference":"Harris, E. H. (1989). The Chlamydomonas sourcebook: a comprehensive guide to biology and laboratory use. Academic Press, San Diego.
","pubmedId":"","doi":"10.1016/C2009-0-02778-0"},{"reference":"Huang, B., Piperno, G., & Luck, D. J. (1979). Paralyzed flagella mutants of Chlamydomonas reinhardtii. Defective for axonemal doublet microtubule arms. J Biol Chem, 254(8), 3091-3099.
","pubmedId":"429335","doi":""},{"reference":"Hutner, S. H., Provasoli, L., Schatz, A., & Haskins, C. P. (1950). Some approaches to the study of the role of metals in the metabolism of microorganisms. Proc Am Philos Soc, 94(2), 152-170.
","pubmedId":"","doi":""},{"reference":"Iwasa, K., & Murakami, S. (1968). Palmelloid formation of Chlamydomonas. Physiologia Plantarum, 21(6), 1224-1233.
","pubmedId":"","doi":"10.1111/j.1399-3054.1968.tb07353.x"},{"reference":"Kamiya, R. (1988). Mutations at twelve independent loci result in absence of outer dynein arms in Chlamydomonas reinhardtii. J Cell Biol, 107(6), 2253-2258.
","pubmedId":"2974040","doi":"10.1083/jcb.107.6.2253"},{"reference":"Kamiya, R., & Yagi, T. (2014). Functional diversity of axonemal dyneins as assessed by in vitro and in vivo motility assays of Chlamydomonas mutants. Zoolog Sci, 31(10), 633-644.
","pubmedId":"25284382","doi":"10.2108/zs140066"},{"reference":"Khona, D. K., Shirolikar, S. M., Gawde, K. K., Hom, E., Deodhar, M. A., & D'Souza, J. S. (2016). Characterization of salt stress-induced palmelloids in the green alga, Chlamydomonas reinhardtii. Algal Res, 16, 434-448.
","pubmedId":"","doi":"10.1016/j.algal.2016.03.035"},{"reference":"King, S. M., Yagi, T., & Kamiya, R. (2023). Chapter 4 - Axonemal dyneins: genetics, structure, and motor activity. In S. K. Dutcher (Ed.), The Chlamydomonas Sourcebook (3rd Edition) (pp. 79-131). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-822508-0.00002-2"},{"reference":"Marshall, W. F. (2024). Chlamydomonas as a model system to study cilia and flagella using genetics, biochemistry, and microscopy. Front Cell Dev Biol, 12, 1412641.
","pubmedId":"38872931","doi":"10.3389/fcell.2024.1412641"},{"reference":"Mitchell, D. R., & Yamamoto, R. (2023). Chapter 5 - Axonemal dynein preassembly. In S. K. Dutcher (Ed.), The Chlamydomonas Sourcebook (3rd Edition) (pp. 133-155). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-822508-0.00013-7"},{"reference":"Mitchison, H. M., Schmidts, M., Loges, N. T., Freshour, J., Dritsoula, A., Hirst, R. A.,…Mitchell, D. R. (2012). Mutations in axonemal dynein assembly factor DNAAF3 cause primary ciliary dyskinesia. Nat Genet, 44(4), 381-389.
","pubmedId":"22387996","doi":"10.1038/ng.1106"},{"reference":"Omran, H., Kobayashi, D., Olbrich, H., Tsukahara, T., Loges, N. T., Hagiwara, H.,…Takeda, H. (2008). Ktu/PF13 is required for cytoplasmic pre-assembly of axonemal dyneins. Nature, 456(7222), 611-616.
","pubmedId":"19052621","doi":"10.1038/nature07471"},{"reference":"Ostrowski, L. E., Dutcher, S. K., & Lo, C. W. (2011). Cilia and models for studying structure and function. Proc Am Thorac Soc, 8(5), 423-429.
","pubmedId":"21926393","doi":"10.1513/pats.201103-027SD"},{"reference":"Sager, R., & Granick, S. (1953). Nutritional studies with Chlamydomonas reinhardi. Ann N Y Acad Sci, 56(5), 831-838.
","pubmedId":"13139273","doi":"10.1111/j.1749-6632.1953.tb30261.x"},{"reference":"Satir, P., & Christensen, S. T. (2007). Overview of structure and function of mammalian cilia. Annu Rev Physiol, 69, 377-400.
","pubmedId":"17009929","doi":"10.1146/annurev.physiol.69.040705.141236"},{"reference":"Tarkar, A., Loges, N. T., Slagle, C. E., Francis, R., Dougherty, G. W., Tamayo, J. V.,…Omran, H. (2013). DYX1C1 is required for axonemal dynein assembly and ciliary motility. Nat Genet, 45(9), 995-1003.
","pubmedId":"23872636","doi":"10.1038/ng.2707"},{"reference":"Vincensini, L., Blisnick, T., & Bastin, P. (2011). 1001 model organisms to study cilia and flagella. Biol Cell, 103(3), 109-130.
","pubmedId":"21275904","doi":"10.1042/BC20100104"},{"reference":"Wingfield, J. L., & Lechtreck, K. F. (2018). Chlamydomonas basal bodies as flagella organizing centers. Cells, 7(7), 79.
","pubmedId":"30018231","doi":"10.3390/cells7070079"},{"reference":"Witman, G. B. (1986). Isolation of Chlamydomonas flagella and flagellar axonemes. Methods Enzymol, 134, 280-290.
","pubmedId":"3821567","doi":"10.1016/0076-6879(86)34096-5"},{"reference":"Yamamoto, R., Hirono, M., & Kamiya, R. (2010). Discrete PIH proteins function in the cytoplasmic preassembly of different subsets of axonemal dyneins. J Cell Biol, 190(1), 65-71.
","pubmedId":"20603327","doi":"10.1083/jcb.201002081"},{"reference":"Yamamoto, R., Hwang, J., Ishikawa, T., Kon, T., & Sale, W. S. (2021). Composition and function of ciliary inner-dynein-arm subunits studied in Chlamydomonas reinhardtii. Cytoskeleton (Hoboken), 78(3), 77-96.
","pubmedId":"33876572","doi":"10.1002/cm.21662"},{"reference":"Yamamoto, R., Obbineni, J. M., Alford, L. M., Ide, T., Owa, M., Hwang, J.,…Dutcher, S. K. (2017). Chlamydomonas DYX1C1/PF23 is essential for axonemal assembly and proper morphology of inner dynein arms. PLoS Genet, 13(9), e1006996.
","pubmedId":"28892495","doi":"10.1371/journal.pgen.1006996"},{"reference":"Zariwala, M. A., Omran, H., & Ferkol, T. W. (2011). The emerging genetics of primary ciliary dyskinesia. Proc Am Thorac Soc, 8(5), 430-433.
","pubmedId":"21926394","doi":"10.1513/pats.201103-023SD Abstract"}],"title":"Culture method for promoting efficient ciliary growth in the Chlamydomonas pf23 mutant defective in dynein preassembly
","reviews":[],"curatorReviews":[]},{"id":"50d10c8b-16e3-45a8-a095-adc361701e40","decision":"accept","abstract":"Chlamydomonas reinhardtii is an ideal model organism for studying the cytoplasmic preassembly of ciliary dyneins. However, under normal liquid-culture conditions, Chlamydomonas preassembly-deficient mutants often show no cilia or a small number of ciliated cells (i.e., low ciliation ratio), which hinders further analysis of both ciliary dyneins and the phenotypes of these mutants. In this brief report, we present a modified culture method for one of the Chlamydomonas preassembly-deficient mutants, pf23. This method enables researchers to obtain enough pf23 cilia for small-scale biochemical, biophysical, structural and phenotypic analyses.
","acknowledgements":"We would like to thank Drs. Kazuo Inaba (University of Tsukuba) and Takashi Ishikawa (Paul Scherrer Institute), and Mr. Yoshio Inomata/Akinobu Toratani/Motoki Nishizaka (Osaka University) for their help and advice regarding the phenotypic analyses of the pf23 mutant. We would also like to thank Dr. Winfield S. Sale (Emory University) for his support and careful reading of the manuscript.
","authors":[{"affiliations":["Osaka University, Osaka, Ōsaka, Japan"],"departments":["Department of Biological Sciences, Graduate School of Science"],"credit":["conceptualization","fundingAcquisition","writing_originalDraft","investigation","methodology"],"email":"yamamoto.ryousuke.sci@osaka-u.ac.jp","firstName":"Ryosuke","lastName":"Yamamoto","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["Osaka University, Osaka, Ōsaka, Japan"],"departments":["Department of Biological Sciences, Graduate School of Science"],"credit":["conceptualization","supervision","fundingAcquisition","writing_reviewEditing"],"email":"takahide.kon@bio.sci.osaka-u.ac.jp","firstName":"Takahide","lastName":"Kon","submittingAuthor":false,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":null,"extendedData":[],"funding":"R.Y is a recipient of JSPS Grant-in-Aid for Scientific Research (C)(JP24K09454), and T.K was a recipient of JSPS Grant-in-Aid for Scientific Research (B)(JP17H03665).
","image":{"url":"https://portal.micropublication.org/uploads/26404e26cf2928816c3be4970ca42abb.png"},"imageCaption":"This is a schematic illustration of the ciliation method for pf23, the Chlamydomonas preassembly-deficient mutant. The panels [(A) - (F)] in this figure are explained in the \"Methods\" section of the main text. In our laboratory, we check the ciliary growth of the pf23 cells transferred to the SG medium every day [panel (E)]. As early as 1 day after transferring the cells, we often observed that they had grown sufficient cilia to perform small-scale experiments. Panel (E) also shows a ciliated pf23 cell using this method as an inset. This figure was created using PowerPoint (Microsoft).
","imageTitle":"Culture method for the Chlamydomonas pf23 mutant with difficulty growing cilia
","methods":"","reagents":"MUTANT | GENOTYPE | DEFECTS | AVAILABLE FROM |
pf23-1 | pf23 | Cytoplasmic preassembly of several ciliary dyneins (Huang et al., 1979; Yamamoto et al., 2017) | Chlamydomonas resource center (https://www.chlamycollection.org/) |
Chlamydomonas is an excellent model organism (Marshall, 2024; Ostrowski et al., 2011; Vincensini et al., 2011) for studying the composition and function of motile cilia, organelles that play vital roles in various important biological processes in eukaryotes, such as fertility and development (Satir & Christensen, 2007). In this regard, this tiny alga serves as an outstanding organism for studying the pathogenesis of primary ciliary dyskinesia (PCD), a human disease caused by deficient ciliary motility (Chodhari et al., 2004; Zariwala et al., 2011). Motile cilia are powered by motor-protein complexes called \"ciliary dyneins\" (Kamiya & Yagi, 2014; King et al., 2023; Yamamoto et al., 2021). The ciliary dyneins are preassembled in the cytoplasm before being transported into cilia (in a sequential process called \"preassembly\")(Desai et al., 2017; Mitchell & Yamamoto, 2023), and defects in this process also cause PCD in humans [e.g. (Mitchison et al., 2012; Omran et al., 2008; Tarkar et al., 2013)]. Across species, many Chlamydomonas preassembly-deficient mutants have also been identified [e.g. (Dean & Mitchell, 2013; Desai et al., 2015; Huang et al., 1979; Kamiya, 1988; Yamamoto et al., 2010)]. However, researchers often face a major challenge when studying the mutants: under the standard liquid tris-acetate-phosphate (TAP) medium condition (Gorman & Levine, 1965; Harris, 1989), these preassembly mutants often grow no or very short cilia, which are insufficient for various experimental analyses. Through testing several culture conditions, we found that, under a specific condition, a classical preassembly mutant, pf23 (Huang et al., 1979), tends to grow cilia more efficiently than under the standard liquid TAP medium condition. In this brief report, we describe a simple culture method that results in efficient ciliary growth in the pf23 mutant and provides enough cilia for small-scale biochemical, biophysical, structural and phenotypic analyses. In our laboratory, we mainly use this method, or modified versions of it, to isolate cilia from pf23.
Methods
The protocol for our new culture method (Figure 1) is described as follows:
[0] (Before the experiment/culture) Make ~ 200 - 500 mL of the liquid SG [Sager and Granick, or minimal (M)] medium [originally reported in (Sager & Granick, 1953), see also (Harris, 1989) and the Chlamydomonas resource center (https://www.chlamycollection.org/) for basic/conventional compositions] in a conical flask. Place a magnetic stirrer in the SG medium, then autoclave the SG medium and cool it to room temperature. There is no need to prepare bubbling equipment for aeration.
[1] Inoculate the target strain (in this case, pf23) onto the ~ 5 - 10 solid TAP plates (Gorman & Levine, 1965; Harris, 1989), each measuring ~ 90 mm, and ensure the Chlamydomonas cells are distributed evenly across the plates (Figure 1A). Incubate the cells at room temperature under appropriate light (e.g. under a desk lamp) for ~ 3 - 7 days to allow them to grow well and turn the plates green (Figure 1B).
[2] Using an inoculation loop, scrape all the cells (cultured for ~ 3 - 7 days) from all the solid TAP plates and transfer them into the ~ 200 - 500 mL of the liquid SG medium (prepared in [0])(Figure 1C). Since all the cells from the ~ 5 - 10 solid TAP plates are in the liquid SG medium, the medium should be green to dark green.
[3] Gently mix the Chlamydomonas culture in the SG medium (Figure 1D) at room temperature under appropriate light (e.g. under a desk lamp) with the magnetic stirrer (placed in the SG medium in [0]). There is no need for bubbling/aeration. The stirring should be moderately strong enough to disrupt the mother cell walls of palmelloid cells (Iwasa & Murakami, 1968; Khona et al., 2016; Wingfield & Lechtreck, 2018) without disrupting the cells themselves.
[4] Keep the Chlamydomonas culture mixed in the SG medium with the stirrer under light. The cells will often grow cilia 1 - 3 days after being transferred to the liquid SG medium. Using conventional light microscopy (e.g. Olympus BX50, at ~ ×200 magnification), check if the cells are ciliated every day after transferring them to the SG medium (Figure 1E). Once the cells are sufficiently ciliated (e.g. ciliation ratio ≧ ~ 70%), deciliate them (Figure 1F) and collect the cilia using the standard procedure (Craige et al., 2013; Witman, 1986).
Note that all the procedures should be aseptic, and contamination of the Chlamydomonas cultures by bacteria or fungi, especially during mixing in the SG medium, should be avoided. With this method, one of the Chlamydomonas preassembly-deficient mutants, pf23, tends to grow cilia efficiently. To demonstrate its effectiveness, we cultured pf23 using this method (Figure 1E) and conducted an empirical evaluation of ciliation for this brief report. The SG medium used was essentially identical to the medium (https://chlamycollection.org/SG.html) listed at the Chlamydomonas resource center, with only minor modifications. 1 day after transferring to the SG medium, the ciliation ratio of pf23 was approximately ≧ ~ 80%, based on a rough visual estimation. In the past, researchers observed that some preassembly-deficient mutants tend to grow cilia at a late culture stage in the nutrient-rich media (including the TAP medium)[e.g. (Huang et al., 1979; Yamamoto et al., 2010)], i.e., when nutrients in the nutrient-rich media decrease from the initial high amounts. We incorporated this observation into this method, establishing an easy way to grow pf23 cilia. Due to the availability of reagents/chemicals as well as historical reasons, compositions of the SG medium somewhat vary between laboratories studying Chlamydomonas, but we believe most of these media would work well for this method. In addition, in this method, other nutrient-poor Chlamydomonas media [e.g. nitrate-as-nitrogen-source (N) medium (Sager & Granick, 1953), see also the Chlamydomonas resource center (https://www.chlamycollection.org/)] could potentially be used as an alternative to the SG medium.
Conclusion
We have long used this method, or modified versions of it [e.g. usage of \"Hutner’s trace elements\" (Hutner et al., 1950) instead of the trace elements], to isolate sufficient amounts of cilia from pf23, the classical Chlamydomonas preassembly-deficient mutant. This method is simple and convenient, can be performed using standard equipment, and is workable in most laboratories studying Chlamydomonas. This method could also be applicable to other Chlamydomonas ciliary and/or preassembly-deficient mutants that grow very short or few cilia under the standard TAP medium condition, such as pf13 (Huang et al., 1979; Omran et al., 2008) and pf22 (Huang et al., 1979; Mitchison et al., 2012).
","references":[{"reference":"Chodhari, R., Mitchison, H. M., & Meeks, M. (2004). Cilia, primary ciliary dyskinesia and molecular genetics. Paediatr Respir Rev, 5(1), 69-76.
","pubmedId":"15222957","doi":"10.1016/j.prrv.2003.09.005"},{"reference":"Craige, B., Brown, J. M., & Witman, G. B. (2013). Isolation of Chlamydomonas flagella. Curr Protoc Cell Biol, Chapter 3 (Unit 3.41), 1-9.
","pubmedId":"23728744","doi":"10.1002/0471143030.cb0341s59"},{"reference":"Dean, A. B., & Mitchell, D. R. (2013). Chlamydomonas ODA10 is a conserved axonemal protein that plays a unique role in outer dynein arm assembly. Mol Biol Cell, 24(23), 3689-3696.
","pubmedId":"24088566","doi":"10.1091/mbc.E13-06-0310"},{"reference":"Desai, P. B., Dean, A. B., & Mitchell, D. R. (2017). Chapter 4 - Cytoplasmic preassembly and trafficking of axonemal dyneins. In S. M. King (Ed.), Dyneins (The Biology of Dynein Motors)( 2 nd Edition) (pp. 140-161). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-809471-6.00004-8"},{"reference":"Desai, P. B., Freshour, J. R., & Mitchell, D. R. (2015). Chlamydomonas axonemal dynein assembly locus ODA8 encodes a conserved flagellar protein needed for cytoplasmic maturation of outer dynein arm complexes. Cytoskeleton (Hoboken), 72(1), 16-28.
","pubmedId":"25558044","doi":"10.1002/cm.21206"},{"reference":"Gorman, D. S., & Levine, R. P. (1965). Cytochrome f and plastocyanin: their sequence in the photosynthetic electron transport chain of Chlamydomonas reinhardi. Proc Natl Acad Sci U S A, 54(6), 1665-1669.
","pubmedId":"4379719","doi":"10.1073/pnas.54.6.1665"},{"reference":"Harris, E. H. (1989). The Chlamydomonas sourcebook: a comprehensive guide to biology and laboratory use. Academic Press, San Diego.
","pubmedId":"","doi":"10.1016/C2009-0-02778-0"},{"reference":"Huang, B., Piperno, G., & Luck, D. J. (1979). Paralyzed flagella mutants of Chlamydomonas reinhardtii. Defective for axonemal doublet microtubule arms. J Biol Chem, 254(8), 3091-3099.
","pubmedId":"429335","doi":""},{"reference":"Hutner, S. H., Provasoli, L., Schatz, A., & Haskins, C. P. (1950). Some approaches to the study of the role of metals in the metabolism of microorganisms. Proc Am Philos Soc, 94(2), 152-170.
","pubmedId":"","doi":""},{"reference":"Iwasa, K., & Murakami, S. (1968). Palmelloid formation of Chlamydomonas. Physiologia Plantarum, 21(6), 1224-1233.
","pubmedId":"","doi":"10.1111/j.1399-3054.1968.tb07353.x"},{"reference":"Kamiya, R. (1988). Mutations at twelve independent loci result in absence of outer dynein arms in Chlamydomonas reinhardtii. J Cell Biol, 107(6), 2253-2258.
","pubmedId":"2974040","doi":"10.1083/jcb.107.6.2253"},{"reference":"Kamiya, R., & Yagi, T. (2014). Functional diversity of axonemal dyneins as assessed by in vitro and in vivo motility assays of Chlamydomonas mutants. Zoolog Sci, 31(10), 633-644.
","pubmedId":"25284382","doi":"10.2108/zs140066"},{"reference":"Khona, D. K., Shirolikar, S. M., Gawde, K. K., Hom, E., Deodhar, M. A., & D'Souza, J. S. (2016). Characterization of salt stress-induced palmelloids in the green alga, Chlamydomonas reinhardtii. Algal Res, 16, 434-448.
","pubmedId":"","doi":"10.1016/j.algal.2016.03.035"},{"reference":"King, S. M., Yagi, T., & Kamiya, R. (2023). Chapter 4 - Axonemal dyneins: genetics, structure, and motor activity. In S. K. Dutcher (Ed.), The Chlamydomonas Sourcebook (3rd Edition) (pp. 79-131). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-822508-0.00002-2"},{"reference":"Marshall, W. F. (2024). Chlamydomonas as a model system to study cilia and flagella using genetics, biochemistry, and microscopy. Front Cell Dev Biol, 12, 1412641.
","pubmedId":"38872931","doi":"10.3389/fcell.2024.1412641"},{"reference":"Mitchell, D. R., & Yamamoto, R. (2023). Chapter 5 - Axonemal dynein preassembly. In S. K. Dutcher (Ed.), The Chlamydomonas Sourcebook (3rd Edition) (pp. 133-155). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-822508-0.00013-7"},{"reference":"Mitchison, H. M., Schmidts, M., Loges, N. T., Freshour, J., Dritsoula, A., Hirst, R. A.,…Mitchell, D. R. (2012). Mutations in axonemal dynein assembly factor DNAAF3 cause primary ciliary dyskinesia. Nat Genet, 44(4), 381-389.
","pubmedId":"22387996","doi":"10.1038/ng.1106"},{"reference":"Omran, H., Kobayashi, D., Olbrich, H., Tsukahara, T., Loges, N. T., Hagiwara, H.,…Takeda, H. (2008). Ktu/PF13 is required for cytoplasmic pre-assembly of axonemal dyneins. Nature, 456(7222), 611-616.
","pubmedId":"19052621","doi":"10.1038/nature07471"},{"reference":"Ostrowski, L. E., Dutcher, S. K., & Lo, C. W. (2011). Cilia and models for studying structure and function. Proc Am Thorac Soc, 8(5), 423-429.
","pubmedId":"21926393","doi":"10.1513/pats.201103-027SD"},{"reference":"Sager, R., & Granick, S. (1953). Nutritional studies with Chlamydomonas reinhardi. Ann N Y Acad Sci, 56(5), 831-838.
","pubmedId":"13139273","doi":"10.1111/j.1749-6632.1953.tb30261.x"},{"reference":"Satir, P., & Christensen, S. T. (2007). Overview of structure and function of mammalian cilia. Annu Rev Physiol, 69, 377-400.
","pubmedId":"17009929","doi":"10.1146/annurev.physiol.69.040705.141236"},{"reference":"Tarkar, A., Loges, N. T., Slagle, C. E., Francis, R., Dougherty, G. W., Tamayo, J. V.,…Omran, H. (2013). DYX1C1 is required for axonemal dynein assembly and ciliary motility. Nat Genet, 45(9), 995-1003.
","pubmedId":"23872636","doi":"10.1038/ng.2707"},{"reference":"Vincensini, L., Blisnick, T., & Bastin, P. (2011). 1001 model organisms to study cilia and flagella. Biol Cell, 103(3), 109-130.
","pubmedId":"21275904","doi":"10.1042/BC20100104"},{"reference":"Wingfield, J. L., & Lechtreck, K. F. (2018). Chlamydomonas basal bodies as flagella organizing centers. Cells, 7(7), 79.
","pubmedId":"30018231","doi":"10.3390/cells7070079"},{"reference":"Witman, G. B. (1986). Isolation of Chlamydomonas flagella and flagellar axonemes. Methods Enzymol, 134, 280-290.
","pubmedId":"3821567","doi":"10.1016/0076-6879(86)34096-5"},{"reference":"Yamamoto, R., Hirono, M., & Kamiya, R. (2010). Discrete PIH proteins function in the cytoplasmic preassembly of different subsets of axonemal dyneins. J Cell Biol, 190(1), 65-71.
","pubmedId":"20603327","doi":"10.1083/jcb.201002081"},{"reference":"Yamamoto, R., Hwang, J., Ishikawa, T., Kon, T., & Sale, W. S. (2021). Composition and function of ciliary inner-dynein-arm subunits studied in Chlamydomonas reinhardtii. Cytoskeleton (Hoboken), 78(3), 77-96.
","pubmedId":"33876572","doi":"10.1002/cm.21662"},{"reference":"Yamamoto, R., Obbineni, J. M., Alford, L. M., Ide, T., Owa, M., Hwang, J.,…Dutcher, S. K. (2017). Chlamydomonas DYX1C1/PF23 is essential for axonemal assembly and proper morphology of inner dynein arms. PLoS Genet, 13(9), e1006996.
","pubmedId":"28892495","doi":"10.1371/journal.pgen.1006996"},{"reference":"Zariwala, M. A., Omran, H., & Ferkol, T. W. (2011). The emerging genetics of primary ciliary dyskinesia. Proc Am Thorac Soc, 8(5), 430-433.
","pubmedId":"21926394","doi":"10.1513/pats.201103-023SD Abstract"}],"title":"Culture method for promoting efficient ciliary growth in the Chlamydomonas pf23 mutant defective in dynein preassembly
","reviews":[{"reviewer":{"displayName":"Gregory J Pazour"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[]},{"id":"30636700-4d2f-485d-870e-50cae634b6bd","decision":"publish","abstract":"Chlamydomonas reinhardtii is an ideal model organism for studying the cytoplasmic preassembly of ciliary dyneins. However, under normal liquid-culture conditions, Chlamydomonas preassembly-deficient mutants often show no cilia or a small number of ciliated cells (i.e., low ciliation ratio), which hinders further analysis of both ciliary dyneins and the phenotypes of these mutants. In this brief report, we present a modified culture method for one of the Chlamydomonas preassembly-deficient mutants, pf23. This method enables researchers to obtain enough pf23 cilia for small-scale biochemical, biophysical, structural and phenotypic analyses.
","acknowledgements":"We would like to thank Drs. Kazuo Inaba (University of Tsukuba) and Takashi Ishikawa (Paul Scherrer Institute), and Mr. Yoshio Inomata/Akinobu Toratani/Motoki Nishizaka (Osaka University) for their help and advice regarding the phenotypic analyses of the pf23 mutant. We would also like to thank Dr. Winfield S. Sale (Emory University) for his support and careful reading of the manuscript.
","authors":[{"affiliations":["Osaka University, Osaka, Ōsaka, Japan"],"departments":["Department of Biological Sciences, Graduate School of Science"],"credit":["conceptualization","fundingAcquisition","writing_originalDraft","investigation","methodology"],"email":"yamamoto.ryousuke.sci@osaka-u.ac.jp","firstName":"Ryosuke","lastName":"Yamamoto","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null},{"affiliations":["Osaka University, Osaka, Ōsaka, Japan"],"departments":["Department of Biological Sciences, Graduate School of Science"],"credit":["conceptualization","supervision","fundingAcquisition","writing_reviewEditing"],"email":"takahide.kon@bio.sci.osaka-u.ac.jp","firstName":"Takahide","lastName":"Kon","submittingAuthor":false,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":null}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"R.Y is a recipient of JSPS Grant-in-Aid for Scientific Research (C)(JP24K09454), and T.K was a recipient of JSPS Grant-in-Aid for Scientific Research (B)(JP17H03665).
","image":{"url":"https://portal.micropublication.org/uploads/26404e26cf2928816c3be4970ca42abb.png"},"imageCaption":"This is a schematic illustration of the ciliation method for pf23, the Chlamydomonas preassembly-deficient mutant. The panels [(A) - (F)] in this figure are explained in the \"Methods\" section of the main text. In our laboratory, we check the ciliary growth of the pf23 cells transferred to the SG medium every day [panel (E)]. As early as 1 day after transferring the cells, we often observed that they had grown sufficient cilia to perform small-scale experiments. Panel (E) also shows a ciliated pf23 cell using this method as an inset. This figure was created using PowerPoint (Microsoft).
","imageTitle":"Culture method for the Chlamydomonas pf23 mutant with difficulty growing cilia
","methods":"","reagents":"MUTANT | GENOTYPE | DEFECTS | AVAILABLE FROM |
pf23-1 | pf23 | Cytoplasmic preassembly of several ciliary dyneins (Huang et al., 1979; Yamamoto et al., 2017) | Chlamydomonas resource center (https://www.chlamycollection.org/) |
Chlamydomonas is an excellent model organism (Marshall, 2024; Ostrowski et al., 2011; Vincensini et al., 2011) for studying the composition and function of motile cilia, organelles that play vital roles in various important biological processes in eukaryotes, such as fertility and development (Satir & Christensen, 2007). In this regard, this tiny alga serves as an outstanding organism for studying the pathogenesis of primary ciliary dyskinesia (PCD), a human disease caused by deficient ciliary motility (Chodhari et al., 2004; Zariwala et al., 2011). Motile cilia are powered by motor-protein complexes called \"ciliary dyneins\" (Kamiya & Yagi, 2014; King et al., 2023; Yamamoto et al., 2021). The ciliary dyneins are preassembled in the cytoplasm before being transported into cilia (in a sequential process called \"preassembly\")(Desai et al., 2017; Mitchell & Yamamoto, 2023), and defects in this process also cause PCD in humans [e.g. (Mitchison et al., 2012; Omran et al., 2008; Tarkar et al., 2013)]. Across species, many Chlamydomonas preassembly-deficient mutants have also been identified [e.g. (Dean & Mitchell, 2013; Desai et al., 2015; Huang et al., 1979; Kamiya, 1988; Yamamoto et al., 2010)]. However, researchers often face a major challenge when studying the mutants: under the standard liquid tris-acetate-phosphate (TAP) medium condition (Gorman & Levine, 1965; Harris, 1989), these preassembly mutants often grow no or very short cilia, which are insufficient for various experimental analyses. Through testing several culture conditions, we found that, under a specific condition, a classical preassembly mutant, pf23 (Huang et al., 1979), tends to grow cilia more efficiently than under the standard liquid TAP medium condition. In this brief report, we describe a simple culture method that results in efficient ciliary growth in the pf23 mutant and provides enough cilia for small-scale biochemical, biophysical, structural and phenotypic analyses. In our laboratory, we mainly use this method, or modified versions of it, to isolate cilia from pf23.
Methods
The protocol for our new culture method (Figure 1) is described as follows:
[0] (Before the experiment/culture) Make ~ 200 - 500 mL of the liquid SG [Sager and Granick, or minimal (M)] medium [originally reported in (Sager & Granick, 1953), see also (Harris, 1989) and the Chlamydomonas resource center (https://www.chlamycollection.org/) for basic/conventional compositions] in a conical flask. Place a magnetic stirrer in the SG medium, then autoclave the SG medium and cool it to room temperature. There is no need to prepare bubbling equipment for aeration.
[1] Inoculate the target strain (in this case, pf23) onto the ~ 5 - 10 solid TAP plates (Gorman & Levine, 1965; Harris, 1989), each measuring ~ 90 mm, and ensure the Chlamydomonas cells are distributed evenly across the plates (Figure 1A). Incubate the cells at room temperature under appropriate light (e.g. under a desk lamp) for ~ 3 - 7 days to allow them to grow well and turn the plates green (Figure 1B).
[2] Using an inoculation loop, scrape all the cells (cultured for ~ 3 - 7 days) from all the solid TAP plates and transfer them into the ~ 200 - 500 mL of the liquid SG medium (prepared in [0])(Figure 1C). Since all the cells from the ~ 5 - 10 solid TAP plates are in the liquid SG medium, the medium should be green to dark green.
[3] Gently mix the Chlamydomonas culture in the SG medium (Figure 1D) at room temperature under appropriate light (e.g. under a desk lamp) with the magnetic stirrer (placed in the SG medium in [0]). There is no need for bubbling/aeration. The stirring should be moderately strong enough to disrupt the mother cell walls of palmelloid cells (Iwasa & Murakami, 1968; Khona et al., 2016; Wingfield & Lechtreck, 2018) without disrupting the cells themselves.
[4] Keep the Chlamydomonas culture mixed in the SG medium with the stirrer under light. The cells will often grow cilia 1 - 3 days after being transferred to the liquid SG medium. Using conventional light microscopy (e.g. Olympus BX50, at ~ ×200 magnification), check if the cells are ciliated every day after transferring them to the SG medium (Figure 1E). Once the cells are sufficiently ciliated (e.g. ciliation ratio ≧ ~ 70%), deciliate them (Figure 1F) and collect the cilia using the standard procedure (Craige et al., 2013; Witman, 1986).
Note that all the procedures should be aseptic, and contamination of the Chlamydomonas cultures by bacteria or fungi, especially during mixing in the SG medium, should be avoided. With this method, one of the Chlamydomonas preassembly-deficient mutants, pf23, tends to grow cilia efficiently. To demonstrate its effectiveness, we cultured pf23 using this method (Figure 1E) and conducted an empirical evaluation of ciliation for this brief report. The SG medium used was essentially identical to the medium (https://chlamycollection.org/SG.html) listed at the Chlamydomonas resource center, with only minor modifications. 1 day after transferring to the SG medium, the ciliation ratio of pf23 was approximately ≧ ~ 80%, based on a rough visual estimation. In the past, researchers observed that some preassembly-deficient mutants tend to grow cilia at a late culture stage in the nutrient-rich media (including the TAP medium)[e.g. (Huang et al., 1979; Yamamoto et al., 2010)], i.e., when nutrients in the nutrient-rich media decrease from the initial high amounts. We incorporated this observation into this method, establishing an easy way to grow pf23 cilia. Due to the availability of reagents/chemicals as well as historical reasons, compositions of the SG medium somewhat vary between laboratories studying Chlamydomonas, but we believe most of these media would work well for this method. In addition, in this method, other nutrient-poor Chlamydomonas media [e.g. nitrate-as-nitrogen-source (N) medium (Sager & Granick, 1953), see also the Chlamydomonas resource center (https://www.chlamycollection.org/)] could potentially be used as an alternative to the SG medium.
Conclusion
We have long used this method, or modified versions of it [e.g. usage of \"Hutner’s trace elements\" (Hutner et al., 1950) instead of the trace elements], to isolate sufficient amounts of cilia from pf23, the classical Chlamydomonas preassembly-deficient mutant. This method is simple and convenient, can be performed using standard equipment, and is workable in most laboratories studying Chlamydomonas. This method could also be applicable to other Chlamydomonas ciliary and/or preassembly-deficient mutants that grow very short or few cilia under the standard TAP medium condition, such as pf13 (Huang et al., 1979; Omran et al., 2008) and pf22 (Huang et al., 1979; Mitchison et al., 2012).
","references":[{"reference":"Chodhari, R., Mitchison, H. M., & Meeks, M. (2004). Cilia, primary ciliary dyskinesia and molecular genetics. Paediatr Respir Rev, 5(1), 69-76.
","pubmedId":"15222957","doi":"10.1016/j.prrv.2003.09.005"},{"reference":"Craige, B., Brown, J. M., & Witman, G. B. (2013). Isolation of Chlamydomonas flagella. Curr Protoc Cell Biol, Chapter 3 (Unit 3.41), 1-9.
","pubmedId":"23728744","doi":"10.1002/0471143030.cb0341s59"},{"reference":"Dean, A. B., & Mitchell, D. R. (2013). Chlamydomonas ODA10 is a conserved axonemal protein that plays a unique role in outer dynein arm assembly. Mol Biol Cell, 24(23), 3689-3696.
","pubmedId":"24088566","doi":"10.1091/mbc.E13-06-0310"},{"reference":"Desai, P. B., Dean, A. B., & Mitchell, D. R. (2017). Chapter 4 - Cytoplasmic preassembly and trafficking of axonemal dyneins. In S. M. King (Ed.), Dyneins (The Biology of Dynein Motors)( 2 nd Edition) (pp. 140-161). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-809471-6.00004-8"},{"reference":"Desai, P. B., Freshour, J. R., & Mitchell, D. R. (2015). Chlamydomonas axonemal dynein assembly locus ODA8 encodes a conserved flagellar protein needed for cytoplasmic maturation of outer dynein arm complexes. Cytoskeleton (Hoboken), 72(1), 16-28.
","pubmedId":"25558044","doi":"10.1002/cm.21206"},{"reference":"Gorman, D. S., & Levine, R. P. (1965). Cytochrome f and plastocyanin: their sequence in the photosynthetic electron transport chain of Chlamydomonas reinhardi. Proc Natl Acad Sci U S A, 54(6), 1665-1669.
","pubmedId":"4379719","doi":"10.1073/pnas.54.6.1665"},{"reference":"Harris, E. H. (1989). The Chlamydomonas sourcebook: a comprehensive guide to biology and laboratory use. Academic Press, San Diego.
","pubmedId":"","doi":"10.1016/C2009-0-02778-0"},{"reference":"Huang, B., Piperno, G., & Luck, D. J. (1979). Paralyzed flagella mutants of Chlamydomonas reinhardtii. Defective for axonemal doublet microtubule arms. J Biol Chem, 254(8), 3091-3099.
","pubmedId":"429335","doi":""},{"reference":"Hutner, S. H., Provasoli, L., Schatz, A., & Haskins, C. P. (1950). Some approaches to the study of the role of metals in the metabolism of microorganisms. Proc Am Philos Soc, 94(2), 152-170.
","pubmedId":"","doi":""},{"reference":"Iwasa, K., & Murakami, S. (1968). Palmelloid formation of Chlamydomonas. Physiologia Plantarum, 21(6), 1224-1233.
","pubmedId":"","doi":"10.1111/j.1399-3054.1968.tb07353.x"},{"reference":"Kamiya, R. (1988). Mutations at twelve independent loci result in absence of outer dynein arms in Chlamydomonas reinhardtii. J Cell Biol, 107(6), 2253-2258.
","pubmedId":"2974040","doi":"10.1083/jcb.107.6.2253"},{"reference":"Kamiya, R., & Yagi, T. (2014). Functional diversity of axonemal dyneins as assessed by in vitro and in vivo motility assays of Chlamydomonas mutants. Zoolog Sci, 31(10), 633-644.
","pubmedId":"25284382","doi":"10.2108/zs140066"},{"reference":"Khona, D. K., Shirolikar, S. M., Gawde, K. K., Hom, E., Deodhar, M. A., & D'Souza, J. S. (2016). Characterization of salt stress-induced palmelloids in the green alga, Chlamydomonas reinhardtii. Algal Res, 16, 434-448.
","pubmedId":"","doi":"10.1016/j.algal.2016.03.035"},{"reference":"King, S. M., Yagi, T., & Kamiya, R. (2023). Chapter 4 - Axonemal dyneins: genetics, structure, and motor activity. In S. K. Dutcher (Ed.), The Chlamydomonas Sourcebook (3rd Edition) (pp. 79-131). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-822508-0.00002-2"},{"reference":"Marshall, W. F. (2024). Chlamydomonas as a model system to study cilia and flagella using genetics, biochemistry, and microscopy. Front Cell Dev Biol, 12, 1412641.
","pubmedId":"38872931","doi":"10.3389/fcell.2024.1412641"},{"reference":"Mitchell, D. R., & Yamamoto, R. (2023). Chapter 5 - Axonemal dynein preassembly. In S. K. Dutcher (Ed.), The Chlamydomonas Sourcebook (3rd Edition) (pp. 133-155). Academic Press.
","pubmedId":"","doi":"10.1016/B978-0-12-822508-0.00013-7"},{"reference":"Mitchison, H. M., Schmidts, M., Loges, N. T., Freshour, J., Dritsoula, A., Hirst, R. A.,…Mitchell, D. R. (2012). Mutations in axonemal dynein assembly factor DNAAF3 cause primary ciliary dyskinesia. Nat Genet, 44(4), 381-389.
","pubmedId":"22387996","doi":"10.1038/ng.1106"},{"reference":"Omran, H., Kobayashi, D., Olbrich, H., Tsukahara, T., Loges, N. T., Hagiwara, H.,…Takeda, H. (2008). Ktu/PF13 is required for cytoplasmic pre-assembly of axonemal dyneins. Nature, 456(7222), 611-616.
","pubmedId":"19052621","doi":"10.1038/nature07471"},{"reference":"Ostrowski, L. E., Dutcher, S. K., & Lo, C. W. (2011). Cilia and models for studying structure and function. Proc Am Thorac Soc, 8(5), 423-429.
","pubmedId":"21926393","doi":"10.1513/pats.201103-027SD"},{"reference":"Sager, R., & Granick, S. (1953). Nutritional studies with Chlamydomonas reinhardi. Ann N Y Acad Sci, 56(5), 831-838.
","pubmedId":"13139273","doi":"10.1111/j.1749-6632.1953.tb30261.x"},{"reference":"Satir, P., & Christensen, S. T. (2007). Overview of structure and function of mammalian cilia. Annu Rev Physiol, 69, 377-400.
","pubmedId":"17009929","doi":"10.1146/annurev.physiol.69.040705.141236"},{"reference":"Tarkar, A., Loges, N. T., Slagle, C. E., Francis, R., Dougherty, G. W., Tamayo, J. V.,…Omran, H. (2013). DYX1C1 is required for axonemal dynein assembly and ciliary motility. Nat Genet, 45(9), 995-1003.
","pubmedId":"23872636","doi":"10.1038/ng.2707"},{"reference":"Vincensini, L., Blisnick, T., & Bastin, P. (2011). 1001 model organisms to study cilia and flagella. Biol Cell, 103(3), 109-130.
","pubmedId":"21275904","doi":"10.1042/BC20100104"},{"reference":"Wingfield, J. L., & Lechtreck, K. F. (2018). Chlamydomonas basal bodies as flagella organizing centers. Cells, 7(7), 79.
","pubmedId":"30018231","doi":"10.3390/cells7070079"},{"reference":"Witman, G. B. (1986). Isolation of Chlamydomonas flagella and flagellar axonemes. Methods Enzymol, 134, 280-290.
","pubmedId":"3821567","doi":"10.1016/0076-6879(86)34096-5"},{"reference":"Yamamoto, R., Hirono, M., & Kamiya, R. (2010). Discrete PIH proteins function in the cytoplasmic preassembly of different subsets of axonemal dyneins. J Cell Biol, 190(1), 65-71.
","pubmedId":"20603327","doi":"10.1083/jcb.201002081"},{"reference":"Yamamoto, R., Hwang, J., Ishikawa, T., Kon, T., & Sale, W. S. (2021). Composition and function of ciliary inner-dynein-arm subunits studied in Chlamydomonas reinhardtii. Cytoskeleton (Hoboken), 78(3), 77-96.
","pubmedId":"33876572","doi":"10.1002/cm.21662"},{"reference":"Yamamoto, R., Obbineni, J. M., Alford, L. M., Ide, T., Owa, M., Hwang, J.,…Dutcher, S. K. (2017). Chlamydomonas DYX1C1/PF23 is essential for axonemal assembly and proper morphology of inner dynein arms. PLoS Genet, 13(9), e1006996.
","pubmedId":"28892495","doi":"10.1371/journal.pgen.1006996"},{"reference":"Zariwala, M. A., Omran, H., & Ferkol, T. W. (2011). The emerging genetics of primary ciliary dyskinesia. Proc Am Thorac Soc, 8(5), 430-433.
","pubmedId":"21926394","doi":"10.1513/pats.201103-023SD Abstract"}],"title":"Culture method for promoting efficient ciliary growth in the Chlamydomonas pf23 mutant defective in dynein preassembly
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