From Mary Porter, U of MN, April 2006

The bop (bypass of paralysis) mutants were isolated by Dutcher et al. in a pf10 background, by selection for restoration of motility.


Dutcher SK, Gibbons W, Inwood WB. A genetic analysis of suppressors of the PF10 mutation in Chlamydomonas reinhardtii. Genetics. 1988 Dec;120(4):965-76. doi: 10.1093/genetics/120.4.965. PMID: 3224813; PMCID: PMC1203588.

Dutcher SK, Power J, Galloway RE, Porter ME. Reappraisal of the genetic map of Chlamydomonas reinhardtii. J Hered. 1991 Jul-Aug;82(4):295-301. doi: 10.1093/oxfordjournals.jhered.a111089. PMID: 1880392.

Lin J, Le TV, Augspurger K, Tritschler D, Bower R, Fu G, Perrone C, O'Toole ET, Mills KV, Dymek E, Smith E, Nicastro D, Porter ME. FAP57/WDR65 targets assembly of a subset of inner arm dyneins and connects to regulatory hubs in cilia. Mol Biol Cell. 2019 Oct 1;30(21):2659-2680. doi: 10.1091/mbc.E19-07-0367. Epub 2019 Sep 4. PMID: 31483737; PMCID: PMC6761771.


  • Locus:
  • ARG7
  • Chromosome:
  • 1

From Claire Remacle, University of Liege, April 2006

Phenotype: dies in the dark

This mutant has a deletion of 0.7 kb in the cob gene and adjacent end of the mitochondrial genome. DNA analysis showed 15.1 kb monomer genomes and also dimers resulting from asymmetric fusions at the deleted end. The mutant is respiratory-deficient, and cannot grow on acetate in the dark. It lacks cyanide-sensitive respiration.


Dorthu MP, Remy S, Michel-Wolwertz MR, Colleaux L, Breyer D, Beckers MC, Englebert S, Duyckaerts C, Sluse FE, Matagne RF (1992) Biochemical, genetic and molecular characterization of new respiratory-deficient mutants in Chlamydomonas reinhardtii. Plant Mol Biol 18:759-772

Colin M, Dorthu MP, Duby F, Remacle C, Dinant M, Wolwertz MR, Duyckaerts C, Sluse F, Matagne RF (1995) Mutations affecting the mitochondrial genes encoding the cytochrome oxidase subunit I and apocytochrome b of Chlamydomonas reinhardtii. Mol Gen Genet 249:179-184


  • Locus:
  • cob
  • Chromosome:
  • mitochondrial

From Emilio Fernandez, University of Cordoba, May 2006, by way of Erik Hom

This mutant, which is deficient in L-amino-oxidase activity, was isolated by Fernández and colleagues. A personal communication from Olivier Vallon indicates that reduced LAO expression is also characteristic of nitrate reductase mutants, so this is not necessarily a mutation in the LAO1 structural gene.


Prieto R, Fernández E (1993) Toxicity of and mutagenesis by chlorate are independent of nitrate reductase activity in Chlamydomonas reinhardtii. Mol Gen Genet 237:429-438

Pérez-Alegre M, Franco AR (1998) Resistance to l-methionine-S-sulfoximine in Chlamydomonas reinhardtii is due to an alteration in a general amino acid transport system. Planta 207:20-26

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

The npq1 mutant has normal levels of violaxanthin but cannot convert violaxanthin to antheraxanthin and zeaxanthin during exposure to excessive light, suggesting a defect in the enzyme violaxanthin de-epoxidase. Anwaruzzaman et al. mapped the NPQ1 locus to linkage group IV, between BLD1 and COX3. At the time, no Chlamydomonas homolog of the plant VDE gene had been identified. Subsequently this gene was found, but it is on chromosome 2, inconsistent with these mapping results.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380

Anwaruzzaman M, Chin BL, Li XP, Lohr M, Martinez DA, Niyogi KK (2004) Genomic analysis of mutants affecting xanthophyll biosynthesis and regulation of photosynthetic light harvesting in Chlamydomonas reinhardtii. Photosynth Res 82:265-276


  • Locus:
  • NPQ1
  • Chromosome:
  • 9

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

The npq2 mutant accumulates zeaxanthin and completely lacks antheraxanthin and violaxanthin under all light conditions. This is consistent with a defect in the enzyme zeaxanthin epoxidase.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380

Govindjee, Seufferheld MJ (2002) Non-photochemical quenching of chlorophyll a fluorescence: early history and characterization of two xanthophyll-cycle mutants of Chlamydomonas reinhardtii. Funct Plant Biol 29:1141-1155

From Krishna Niyogi, UC Berkeley, July 2006. This strain replaces CC-3988.

Phenotype: carotenoid deficient; requires arginine

The npq2-2 mutant was isolated as a suppressor of the high-light bleaching phenotype of the parent strain npq1 lor1 arg7-1. Mutants at the NPQ2 locus are deficient in zeaxanthin epoxidase activity. For more information on the NPQ1 and NPQ2 loci, please see Niyogi et al. (1997). For more on lor1, please see CC-2420.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380

Baroli I, Niyogi KK (2000) Molecular genetics of xanthophyll-dependent photoprotection in green algae and plants. Philos Trans R Soc Lond B Biol Sci 355:1385-1393

Polle JE, Niyogi KK, Melis A (2001) Absence of lutein, violaxanthin and neoxanthin affects the functional chlorophyll antenna size of photosystem-II but not that of photosystem-I in the green alga Chlamydomonas reinhardtii. Plant Cell Physiol 42:482-491


  • Locus:
  • NPQ1
  • Chromosome:
  • 2

From Krishna Niyogi, UC Berkeley, July 2006. This strain replaces CC-3989.

Phenotype: carotenoid deficient; requires arginine

The npq2-3 mutant was isolated as a suppressor of the high-light bleaching phenotype of the parent strain npq1 lor1 arg7-8. Mutants at the NPQ2 locus are deficient in zeaxanthin epoxidase activity. For more information on the NPQ1 and NPQ2 loci, please see Niyogi et al. (1997). For more on lor1, please see CC-2420.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380

Baroli I, Niyogi KK (2000) Molecular genetics of xanthophyll-dependent photoprotection in green algae and plants. Philos Trans R Soc Lond B Biol Sci 355:1385-1393

Polle JE, Niyogi KK, Melis A (2001) Absence of lutein, violaxanthin and neoxanthin affects the functional chlorophyll antenna size of photosystem-II but not that of photosystem-I in the green alga Chlamydomonas reinhardtii. Plant Cell Physiol 42:482-491


  • Locus:
  • NPQ1
  • Chromosome:
  • 2

From Krishna Niyogi, UC Berkeley, July 2006. This strain replaces CC-3990.

Phenotype: carotenoid deficient; requires arginine

The npq2-4 mutant was isolated as a suppressor of the high-light bleaching phenotype of the parent strain npq1 lor1 arg7-8. Mutants at the NPQ2 locus are deficient in zeaxanthin epoxidase activity. For more information on the NPQ1 and NPQ2 loci, please see Niyogi et al. (1997). For more on lor1, please see CC-2420.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380

Baroli I, Niyogi KK (2000) Molecular genetics of xanthophyll-dependent photoprotection in green algae and plants. Philos Trans R Soc Lond B Biol Sci 355:1385-1393

Polle JE, Niyogi KK, Melis A (2001) Absence of lutein, violaxanthin and neoxanthin affects the functional chlorophyll antenna size of photosystem-II but not that of photosystem-I in the green alga Chlamydomonas reinhardtii. Plant Cell Physiol 42:482-491


  • Locus:
  • NPQ1
  • Chromosome:
  • 2

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient; requires arginine

The npq1 mutant has normal levels of violaxanthin but cannot convert violaxanthin to antheraxanthin and zeaxanthin during exposure to excessive light, suggesting a defect in the enzyme violaxanthin de-epoxidase. For more on lor1, please see CC-2420.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380


  • Locus:
  • NPQ1
  • Chromosome:
  • 2

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient; requires arginine

The npq1 mutant has normal levels of violaxanthin but cannot convert violaxanthin to antheraxanthin and zeaxanthin during exposure to excessive light, suggesting a defect in the enzyme violaxanthin de-epoxidase. For more on lor1, please see CC-2420.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380


  • Locus:
  • NPQ1
  • Chromosome:
  • 2

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

Mutants at the NPQ2 locus are deficient in zeaxanthin epoxidase activity (see Niyogi et al., 1997). For more on lor1, please see CC-2420.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380

Baroli I, Niyogi KK (2000) Molecular genetics of xanthophyll-dependent photoprotection in green algae and plants. Philos Trans R Soc Lond B Biol Sci 355:1385-1393

Polle JE, Niyogi KK, Melis A (2001) Absence of lutein, violaxanthin and neoxanthin affects the functional chlorophyll antenna size of photosystem-II but not that of photosystem-I in the green alga Chlamydomonas reinhardtii. Plant Cell Physiol 42:482-491

Holub O, Seufferheld MJ, Gohlke C, Govindjee, Heiss GJ, Clegg RM (2007) Fluorescence lifetime imaging microscopy of Chlamydomonas reinhardtii: non-photochemical quenching mutants and the effect of photosynthetic inhibitors on the slow chlorophyll fluorescence transient. J Microsc 226(Pt 2):90-120

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

The npq1 mutant has normal levels of violaxanthin but cannot convert violaxanthin to antheraxanthin and zeaxanthin during exposure to excessive light, suggesting a defect in the enzyme violaxanthin de-epoxidase. For more on lor1, please see CC-2420.


Niyogi KK, Bjorkman O, Grossman AR (1997) Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching. Plant Cell 9:1369-1380


  • Locus:
  • NPQ1
  • Chromosome:
  • 2

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

This is a carotenoid-deficient mutant, pale green to nearly white in color, that was shown by McCarthy et al. to be deficient in the PSY1 gene encoding phytoene synthase. Other mutations at the same locus, previously identified as LTS1 by Chemerilova (1978), include w-7 (CC-2843), lts1-135 (CC-1944), and FN68 (CC-2682). The lts1-203 mutation is an Ala to Asp change at amino acid residue 211 in the PYS1 gene.


Chemerilova VI (1978) Study of pigmentation-modifying mutations in Chlamydomonas reinhardi strains of different ploidy. II. Compounds forlts1 mutations and their use for obtaining triploid cultures. Genetika 14:154-162

McCarthy SS, Kobayashi MC, Niyogi KK (2004) White mutants of Chlamydomonas reinhardtii are defective in phytoene synthase. Genetics 168:1249-1257


  • Locus:
  • LTS1 [PSY1]
  • Chromosome:
  • 11

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

This is a carotenoid-deficient, light-sensitive mutant. Please see CC-4109 for more information on the LTS1 [PSY1] locus.


McCarthy SS, Kobayashi MC, Niyogi KK (2004) White mutants of Chlamydomonas reinhardtii are defective in phytoene synthase. Genetics 168:1249-1257


  • Locus:
  • LTS1 [PSY1]
  • Chromosome:
  • 11

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

This is a carotenoid-deficient, light-sensitive mutant. Please see CC-4109 for more information on the LTS1 [PSY1] locus. The lts1-206 mutation changes Trp332 of the PSY1 gene to a stop.


McCarthy SS, Kobayashi MC, Niyogi KK (2004) White mutants of Chlamydomonas reinhardtii are defective in phytoene synthase. Genetics 168:1249-1257


  • Locus:
  • LTS1 [PSY1]
  • Chromosome:
  • 11

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

This is a carotenoid-deficient, light-sensitive mutant. Please see CC-4109 for more information on the LTS1 [PSY1] locus. The lts1-207 mutation changes Gln10 of the PSY1 gene to a stop.


McCarthy SS, Kobayashi MC, Niyogi KK (2004) White mutants of Chlamydomonas reinhardtii are defective in phytoene synthase. Genetics 168:1249-1257


  • Locus:
  • LTS1 [PSY1]
  • Chromosome:
  • 11

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

This is a carotenoid-deficient, light-sensitive mutant. Please see CC-4109 for more information on the LTS1 [PSY1] locus.


McCarthy SS, Kobayashi MC, Niyogi KK (2004) White mutants of Chlamydomonas reinhardtii are defective in phytoene synthase. Genetics 168:1249-1257


  • Locus:
  • LTS1 [PSY1]
  • Chromosome:
  • 11

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

This is a carotenoid-deficient, light-sensitive mutant. Please see CC-4109 for more information on the LTS1 [PSY1] locus.


McCarthy SS, Kobayashi MC, Niyogi KK (2004) White mutants of Chlamydomonas reinhardtii are defective in phytoene synthase. Genetics 168:1249-1257


  • Locus:
  • LTS1 [PSY1]
  • Chromosome:
  • 11

From Krishna Niyogi, UC Berkeley, July 2006

Phenotype: carotenoid deficient

This is a carotenoid-deficient, light-sensitive mutant. Please see CC-4109 for more information on the LTS1 [PSY1] locus.


McCarthy SS, Kobayashi MC, Niyogi KK (2004) White mutants of Chlamydomonas reinhardtii are defective in phytoene synthase. Genetics 168:1249-1257


  • Locus:
  • LTS1 [PSY1]
  • Chromosome:
  • 11

From Peter Luykx, University of Miami, 2006, his PL69

Phenotype: deficient in osmoregulation

Please see CC-4017 for more information on the osm13 mutant. This strain was received as “nit1 nit2?”. Since the original mutant was a NIT1 insertion, we are uncertain how this strain was derived, and what its nit status is. This strain should be maintained on TAP medium supplemented with 0.06M sucrose or lactose.


  • Locus:
  • NIT1
  • Chromosome:
  • 9

From Peter Luykx, University of Miami, 2006, his PL138

Phenotype: deficient in osmoregulation

This is the osm1 mutant in a nit1-305 background. Please see CC-3699 for more information on this mutant. This strain should be maintained on TAP medium supplemented with 0.06M sucrose or lactose. This strain has the wild-type allele at the NIT2 locus.


  • Locus:
  • NIT1
  • Chromosome:
  • 9

From Peter Luykx, University of Miami, 2006, his PL156

Phenotype: deficient in osmoregulation

This is the osm17 mutant in a nit1 background. Please see CC-4023 for more information on this mutant. This strain should be maintained on TAP medium supplemented with 0.06M sucrose or lactose. This strain has the wild-type allele at the NIT2 locus.


  • Locus:
  • NIT1
  • Chromosome:
  • 9

From Peter Luykx, University of Miami, 2006, his PL157.

Phenotype: deficient in osmoregulation

This is the osm17 mutant in a nit1 background. Please see CC-4023 for more information on this mutant. This strain should be maintained on TAP medium supplemented with 0.06M sucrose or lactose. This strain has the wild-type allele at the NIT2 locus.


  • Locus:
  • NIT1
  • Chromosome:
  • 9

From Patrice Hamel, Ohio State University, November 2006

Phenotype: requires acetate

The ccs1 mutants are alleles at a locus originally defined by the ac206 mutation (see CC-939 for more information on that mutant). The protein encoded by this locus is required for synthesis of cytochrome c.


Xie Z, Merchant S (1998) A novel pathway for cytochromes c biogenesis in chloroplasts. Biochim Biophys Acta 1365:309-318

Dreyfuss BW, Hamel PP, Nakamoto SS, Merchant S (2003) Functional analysis of a divergent system II protein, Ccs1, involved in c-type cytochrome biogenesis. J Biol Chem 278:2604-2613


  • Locus:
  • CCS1
  • Chromosome:
  • 13

From Patrice Hamel, Ohio State University, November 2006

Phenotype: requires acetate

The ccs1 mutants are alleles at a locus originally defined by the ac206 mutation (see CC-939 for more information on that mutant). The protein encoded by this locus is required for synthesis of cytochrome c.


Xie Z, Merchant S (1998) A novel pathway for cytochromes c biogenesis in chloroplasts. Biochim Biophys Acta 1365:309-318

Dreyfuss BW, Hamel PP, Nakamoto SS, Merchant S (2003) Functional analysis of a divergent system II protein, Ccs1, involved in c-type cytochrome biogenesis. J Biol Chem 278:2604-2613


  • Locus:
  • CCS1
  • Chromosome:
  • 13

From Patrice Hamel, Ohio State University, November 2006

Phenotype: requires acetate

This is an insertional mutant in the PCY1 or PETE gene encoding plastocyanin. For more information on this locus, please see the following papers:


Li HH, Quinn J, Culler D, Girard-Bascou J, Merchant S (1996) Molecular genetic analysis of plastocyanin biosynthesis in Chlamydomonas reinhardtii. J Biol Chem 271:31283-31289

Eriksson M, Moseley JL, Tottey S, Del Campo JA, Quinn J, Kim Y, Merchant S (2004) Genetic dissection of nutritional copper signaling in chlamydomonas distinguishes regulatory and target genes. Genetics 168:795-807


  • Locus:
  • PCY1 [PETE]
  • Chromosome:
  • 3

From Patrice Hamel, Ohio State University, November 2006

Phenotype: requires acetate

This is an insertional mutation in the CCS5 gene.


Gabilly ST, Dreyfuss BW, Karamoko M, Corvest V, Kropat J, Page MD, Merchant SS, Hamel PP (2010) CCS5, a thioredoxin-like protein involved in the assembly of plastid c-type cytochromes. J Biol Chem 285:29738-29749


  • Locus:
  • CCS5
  • Chromosome:
  • 9

From Patrice Hamel, Ohio State University, November 2006

Phenotype: requires acetate

This is an insertional mutation in the CCS5 gene.


Gabilly ST, Dreyfuss BW, Karamoko M, Corvest V, Kropat J, Page MD, Merchant SS, Hamel PP (2010) CCS5, a thioredoxin-like protein involved in the assembly of plastid c-type cytochromes. J Biol Chem 285:29738-29749


  • Locus:
  • CCS5
  • Chromosome:
  • 9

From Patrice Hamel, Ohio State University, November 2006

Phenotype: requires acetate

This is an insertional mutation in the CCS5 gene.


Gabilly ST, Dreyfuss BW, Karamoko M, Corvest V, Kropat J, Page MD, Merchant SS, Hamel PP (2010) CCS5, a thioredoxin-like protein involved in the assembly of plastid c-type cytochromes. J Biol Chem 285:29738-29749


  • Locus:
  • CCS5
  • Chromosome:
  • 9

From Carolyn Silflow, University of Minnesota, January 2007

Phenotype: variable number of flagella

Strain 5E8I4F(vfl1-2::NIT1) mt- is an insertional allele of vfl1 obtained by transformation with the NIT1 gene.

For more information on the VFL1 locus, please see CC-1388.


Tam LW, Lefebvre PA (1993) Cloning of flagellar genes in Chlamydomonas reinhardtii by DNA insertional mutagenesis. Genetics 135:375-384

Silflow CD, LaVoie M, Tam LW, Tousey S, Sanders M, Wu W, Borodovsky M, Lefebvre PA (2001) The Vfl1 Protein in Chlamydomonas localizes in a rotationally asymmetric pattern at the distal ends of the basal bodies. J Cell Biol 153:63-74


  • Locus:
  • VFL1
  • Chromosome:
  • 8