From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac115 and ac115a mutations were UV-induced in Levine’s laboratory.

Kuchka et al. determined that the AC115 gene product specifically promotes translation of the mRNA from the chloroplast psbD gene, encoding the photosystem II D2 protein. In ac115 mutants other proteins of the PS II complex are synthesized but cannot assemble into a stable complex, and are post-translationally degraded.

However, there is a problem: the gene sequenced by Kuchka and collaborators (accessions AF045466 and AF045467; Rattanachaikunsopon et al., 1999) turns out to be on chromosome 8 in the JGI v4 genome sequence, whereas the mutation was mapped to linkage group (chromosome) 1. This conflict has not yet been resolved.


Kuchka MR, Mayfield SP, Rochaix JD (1988) Nuclear mutations specifically affect the synthesis and/or degradation of the chloroplast-encoded D2 polypeptide of photosystem II in Chlamydomonas reinhardtii. EMBO J 7:319-24

Wu HY, Kuchka MR (1995) A nuclear suppressor overcomes defects in the synthesis of the chloroplast psbD gene product caused by mutations in two distinct nuclear genes of Chlamydomonas. Curr Genet 27:263-9

Rattanachaikunsopon P, Rosch C, Kuchka MR (1999) Cloning and characterization of the nuclear AC115 gene of Chlamydomonas reinhardtii. Plant Mol Biol 39:1-10


  • Locus:
  • AC115
  • Chromosome:
  • 1,8

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac141 mutation was UV-induced in Levine’s laboratory. It is easy to score in crosses, and mutant clones form small colonies on tetrad plates.

This mutant was originally reported to be deficient in cytochrome b559 and quinone [Q] activities and had altered thylakoid membranes, with more single discs than wild type strains. Later reports grouped it with other mutants with pleiotropic deficiencies in PS II polypeptides.

A suppressor mutation that restored photosynthetic competence appeared spontaneously in the original ac141 stock and was mapped to linkage group XI by Cosbey et al.,. This mutation also suppressed arg1, ac80a, and ac115, but not arg2, ac31a, ac14k, ac28, ac20, or ac21 The stock containing the original suppressor mutation appears to have been lost.


Levine RP and Volkmann D (1961) Mutants with impaired photosynthesis in Chlamydomonas reinhardi. Biochem Biophys Res Commun. 6:264-269

Hastings PJ, Levine EE, Cosbey E, Hudock MO, Gillham NW, Surzycki SJ, Loppes R, Levine RP (1965) The Linkage groups of Chlamydomonas reinhardi. Microb Genet Bull 23:20-21

Levine RP, Gorman DS (1966) Photosynthetic electron transport chain of Chlamydomonas reinhardi. 3. Light-induced absorbance changes in chloroplast fragments of the wild type and mutant strains. Plant Physiol 41:1293-300

Teichler-Zallen D, Levine RP (1967) An Electron Spin Resonance Study of Manganese in Wild-Type and Mutant Strains of Chlamydomonas reinhardi. Plant Physiol 42:1643-7

Lavorel J, Levine RP (1968) Fluorescence Properties of Wild-Type Chlamydomonas reinhardi and Three Mutant Strains Having Impaired Photosynthesis. Plant Physiol 43:1049-55

Goodenough UW, Levine RP (1969) Chloroplast Ultrastructure in Mutant Strains of Chlamydomonas reinhardi Lacking Components of the Photosynthetic Apparatus. Plant Physiol 44:990-1000

Chua NH, Levine RP (1969) The photosynthetic electron transport chain of Chlamydomonas reinhardi. 8. The 520 nm light-induced absorbance change in the wild-type and mutant strains. Plant Physiol 44:1-6


  • Locus:
  • AC141
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

Please see CC-549 for more information on the ac141 mutation.


  • Locus:
  • AC141
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac177 mutation was UV-induced in Levine’s laboratory. Levine and Goodenough (1970) listed this mutant as nonphotosynthetic; the specific defect has not been not determined.

This locus is erroneously labeled ac17 rather than ac177 on the linkage group XIV map in Kathir et al. (2003).

This strain was easy to score at its most recent check, a good marker for this linkage group.


Levine RP, Goodenough UW (1970) The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Annu Rev Genet 4:397-408

Kathir P, LaVoie M, Brazelton WJ, Haas NA, Lefebvre PA, Silflow CD (2003) Molecular map of the Chlamydomonas reinhardtii nuclear genome. Eukaryot Cell 2:362-79


  • Locus:
  • AC177
  • Chromosome:
  • 13

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This strain is scorable, but CC-551 ac177 mt+ may be a better choice for crosses. Please see that strain for more information on the AC177 locus.


  • Locus:
  • AC177
  • Chromosome:
  • 13

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac200 mutation was induced in Levine’s laboratory with nitrous acid, but no genetic analysis was done there, and nothing was published on it. A cross in 1982 suggested possible linkage to group III, but this has not been pursued further. The acetate requirement was scorable at last check.


  • Locus:
  • AC200
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac207 mutation was UV-induced in Levine’s laboratory. Levine and Goodenough listed the ac207 mutant as nonphotosynthetic. The cause of the defect has not been determined.

This strain has a somewhat leaky but scorable phenotype.


Levine RP, Goodenough UW (1970) The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Annu Rev Genet 4:397-408


  • Locus:
  • AC207
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate on copper-containing medium.

Please see CC-556 for more information on the ac208 mutation.

The CC-555 strain has given a high degree of lethality and aberrant segregations in past crosses. CC-556 ac208 mt- or CC-1729 ac208 mt- would probably be better choices for genetic analysis if mating type is not an issue.


Hill KL, Merchant S (1992) In Vivo Competition between Plastocyanin and a Copper-Dependent Regulator of the Chlamydomonas reinhardtii Cytochrome c(6) Gene. Plant Physiol 100:319-26

Quinn J, Li HH, Singer J, Morimoto B, Mets L, Kindle K, Merchant S (1993) The plastocyanin-deficient phenotype of Chlamydomonas reinhardtii Ac-208 results from a frame-shift mutation in the nuclear gene encoding preapoplastocyanin. J Biol Chem 15:7832-41


  • Locus:
  • AC208 [PETE]
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate on copper-containing medium.

The ac208 mutation was UV-induced in a germinating zygote in Levine’s laboratory.

This mutant is acetate-requiring on copper-containing medium. Copper deprivation induces expression of cytochrome c6, which bypasses the requirement for acetate. This mutant has been extensively studied by Merchant and colleagues. Quinn et al. showed that the alteration is a frameshift mutation in the gene encoding plastocyanin.

We have two stocks of this mutant in mating type minus, this one and CC-1729 in recent years we have generally sent out CC-1729, although both are listed in the core collection.


Hill KL, Merchant S (1992) In Vivo Competition between Plastocyanin and a Copper-Dependent Regulator of the Chlamydomonas reinhardtii Cytochrome c(6) Gene. Plant Physiol 100:319-26

Quinn J, Li HH, Singer J, Morimoto B, Mets L, Kindle K, Merchant S (1993) The plastocyanin-deficient phenotype of Chlamydomonas reinhardtii Ac-208 results from a frame-shift mutation in the nuclear gene encoding preapoplastocyanin. J Biol Chem 15:7832-41


  • Locus:
  • AC208 [PETE]
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: impaired motility

McVittie reported that the PF7 and PF8 loci were linked, and Ramanis and Luck mapped them to linkage group XIX. Neither mutant crosses well, however, and they haven’t received much attention beyond these early crosses.


McVittie A (1972) Flagellum mutants of Chlamydomonas reinhardii. J Gen Microbiol 71:25-40

McVittie A (1972) Genetic studies on flagellum mutants of Chlamydomonas reinhardtii. Genet Res 19:157-164

Ramanis Z, Luck DJ (1986) Loci affecting flagellar assembly and function map to an unusual linkage group in Chlamydomonas reinhardtii. Proc Natl Acad Sci U S A 83:423-6


  • Locus:
  • PF8
  • Chromosome:
  • 17

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac10 isolate was a tetrad product from a cross in Levine’s lab involving the original ac9 mutant but for some years was given a separate designation. It maps to the same location as ac9 on linkage group XII/XIII but has not been analyzed further. For more information on ac9, please see CC-521.


  • Locus:
  • AC9
  • Chromosome:
  • 12

From Ursula Goodenough, Harvard University, May 1978

Phenotype: somewhat chlorophyll deficient

The ac55 and ac55a mutants were UV-induced in Levine’s laboratory, and are yellow to yellow-green in color on all media. This strain is noticeably yellow-green, and is easy to score in crosses.


Levine RP and Volkmann D (1961) Mutants with impaired photosynthesis in Chlamydomonas reinhardi. Biochem Biophys Res Commun 6:264-269

Levine RP, Goodenough UW (1970) The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Annu Rev Genet 4:397-408


  • Locus:
  • AC55
  • Chromosome:
  • 4

From Ursula Goodenough, Harvard University, May 1978

Phenotype: somewhat chlorophyll deficient

The ac55 and ac55a mutants were UV-induced in Levine’s laboratory, and are yellow to yellow-green in color on all media. This strain is noticeably yellow-green, and is easy to score in crosses.


Levine RP and Volkmann D (1961) Mutants with impaired photosynthesis in Chlamydomonas reinhardi. Biochem Biophys Res Commun 6:264-269

Levine RP, Goodenough UW (1970) The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Annu Rev Genet 4:397-408


  • Locus:
  • AC55
  • Chromosome:
  • 4

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac210 mutation was UV-induced and isolated in a germinating zygote in Levine’s laboratory.

This is not an easy mutant to score. The nearby locus AC207 may be a better choice for the right arm of linkage group III.


Levine RP, Goodenough UW (1970) The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Annu Rev Genet 4:397-408


  • Locus:
  • AC210
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: impaired motility

McVittie reported that the PF7 and PF8 loci were linked, and Ramanis and Luck mapped them to linkage group XIX. Neither mutant crosses well, however, and they haven’t received much attention beyond these early crosses. formation.


McVittie A (1972) Flagellum mutants of Chlamydomonas reinhardii. J Gen Microbiol 71:525-40

McVittie A (1972) Genetic studies on flagellum mutants of Chlamydomonas reinhardtii. Genet Res. 19:157-164

Ramanis Z, Luck DJ (1986) Loci affecting flagellar assembly and function map to an unusual linkage group in Chlamydomonas reinhardtii. Proc Natl Acad Sci U S A 83:423-6


  • Locus:
  • PF7
  • Chromosome:
  • 17

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This mutant was isolated by Eversole, and originally called EV1a. The ac1 stock that was preserved by Levine’s laboratory was the result of cloning for large colony size and crossing the selected clone back to a wild type strain.

This strain has been easy to score in crosses.


Eversole RA (1956) Biochemical Mutants of Chlamydomonas reinhardi. Amer J Bot 43:404-407

Ebersold WT, Levine RP, Levine EE, Olmsted MA (1962) Linkage maps in Chlamydomonas reinhardi. Genetics 47:531-543

Goodenough UW and Levine RP (1969) Chloroplast Ultrastructure in Mutant Strains of Chlamydomonas reinhardi Lacking Components of the Photosynthetic Apparatus. Plant Physiol 44:990-1000

Epel BL and Butler WL (1972) A spectroscopic analysis of a high fluorescent mutant of Chlamydomonas reinhardi. Biophys J 12:922-929


  • Locus:
  • AC1
  • Chromosome:
  • 7

Hurley Shepherd and Edwin Wurtz, Boynton-Gillham laboratory, Duke University

Phenotype: requires acetate

This is a nuclear mutant isolated in a mutagenesis experiment with bromodeoxyuridine. It was cleanly acetate requiring at last check, but has not been characterized further.

From Nam-Hai Chua, Rockefeller University, June 1978

Phenotype: requires acetate

This stock is probably a double or even triple mutant strain, containing F34 and one or more unidentified mutations, and is not recommended for use but is being retained for historical reasons. CC-2340 is a preferred alternative.


Girard-Bascou J, Pierre Y, Drapier D (1992) A nuclear mutation affects the synthesis of the chloroplast psbA gene production Chlamydomonas reinhardtii. Curr Genet 22:47-52

From Kathleen Peterson and W.T. Ebersold, UCLA, June 1978

Phenotype: inhibitor resistant (colchicine)

This colchicine-resistant strain has not been fully characterized, but has been retained in the collection in hopes that it will receive further study.

From Kathleen Peterson and W.T. Ebersold, UCLA, June 1978

Phenotype: requires acetate

This is a good genetic marker, cleanly acetate-requiring and easy to score.

Levine and Goodenough (1970) listed the ac28 mutant among those having normal pigment and CO2 fixation; the reason for the acetate requirement is unknown. At least nine allelic mutants were isolated in Levine’s laboratory, of which only this one, formerly ac28d, is extant.


Levine RP and Goodenough UW (1970) The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Annu Rev Genet 4:397-408

Ebersold WT, Levine RP, Levine EE, Olmsted MA (1962) Linkage maps in Chlamydomonas reinhardi. Genetics 47:531-543


  • Locus:
  • AC28
  • Chromosome:
  • 3

From Kathleen Peterson and W.T. Ebersold, UCLA, June 1978

Phenotype: requires nicotinamide

The nic1 mutation was obtained by Eversole, and is one of the few genetic markers for linkage group XV. This strain has given relatively high lethality in some crosses, but is easy to score by sensitivity to 3-acetyl pyridine.

Lin et al. demonstrated that this is a mutation in the gene encoding nicotinate-/nicotinamide-nucleotide adenylyltransferase.


Eversole RA (1956) Biochemical Mutants of Chlamydomonas reinhardi. Amer J Bot 43:404-407

Lin H, Kwan AL, Dutcher SK (2010) Synthesizing and salvaging NAD: lessons learned from Chlamydomonas reinhardtii. PLoS Genet 6


  • Locus:
  • NIC1
  • Chromosome:
  • 14

From Kathleen Peterson and W.T. Ebersold, UCLA, June 1978

Phenotype: requires nicotinamide

The nic1 mutation was obtained by Eversole, and is one of the few genetic markers for linkage group XV. This strain has given relatively high lethality in some crosses, but is easy to score by sensitivity to 3-acetyl pyridine.

Lin et al. demonstrated that this is a mutation in the gene encoding nicotinate-/nicotinamide-nucleotide adenylyltransferase.


  • Locus:
  • PAB1
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, June 1978

Phenotype: impaired motility

The pf1 mutant has a scorable motility defect, and is one of relatively few genetic markers on linkage group V. For research on motility, the better-characterized CC-1024 is preferred (please see that strain for more information). However, CC-602 is widely used in a student laboratory exercise in combination with CC-1032 pf14 mt+, to demonstrate complementation of flagellar mutations in newly formed zygotes (see the mating and dikaryon rescue kit)

This is probably the pf1 allele isolated by Lewin and subsequently mapped in Levine’s laboratory.


  • Locus:
  • PF1 [RSP4]
  • Chromosome:
  • 5

From Ursula Goodenough, Harvard University, June 1978

Phenotype: impaired motility

This mutant wiggles or swims slowly, and is easy to score. The flagella show shallow bend forms. CC-1026 is preferred for research studies, since it is better characterized, but CC-604 does show the correct linkage position for this mutation and is an acceptable mt- stock of pf3 for genetic analysis. Please see CC-1026 for more information on this locus.


  • Locus:
  • PF3
  • Chromosome:
  • 8

From Ursula Goodenough, Harvard University, June 1978

Phenotype: impaired motility

This mutant is easy to score, well-behaved in crosses, and a good marker for linkage group II.

Please see CC-1031 for more information on the PF12 locus.


  • Locus:
  • PF12
  • Chromosome:
  • 2

From Ursula Goodenough, Harvard University, June 1978

Phenotype: impaired motility

This mutant is easy to score, and is an excellent marker for the right arm of linkage group VI.

The pf14 mutants lack radial spokes and are missing a set of 17 spoke-associated polypeptides. Huang and colleagues in Luck’s laboratory used dikaryon and revertant analysis to identify one of these as a probable gene product. The gene was subsequently cloned and sequenced in Rosenbaum’s laboratory, confirming identity of the gene product as radial spoke protein 3.


Huang B, Rifkin MR, Luck DJ (1977) Temperature-sensitive mutations affecting flagellar assembly and function in Chlamydomonas reinhardtii. J Cell Biol 72:67-85

Luck DJ, Piperno G, Ramanis Z, Huang B (1977) Flagellar mutants of Chlamydomonas: studies of radial spoke-defective strains by dikaryon and revertant analysis. Proc Natl Acad Sci USA 74:3456-3460

Diener DR, Ang LH, Rosenbaum JL (1993) Assembly of flagellar radial spoke proteins in Chlamydomonas: identification of the axoneme binding domain of radial spoke protein 3. J Cell Biol 123:183-190

Yang P, Diener DR, Yang C, Kohno T, Pazour GJ, Dienes JM, Agrin NS, King SM, Sale WS, Kamiya R, Rosenbaum JL, Witman GB (2006) Radial spoke proteins of Chlamydomonas flagella. J Cell Sci 119(Pt 6):1165-1174


  • Locus:
  • PF14 [RSP3]
  • Chromosome:
  • 6

From Ursula Goodenough, Harvard University, June 1978

CC-620 and CC-621 are subclones of the Smith 137c wild type strain used in R.P. Levine’s laboratory at Harvard in the 1960s, selected for high mating efficiency. They are often used as demonstration material for Chlamydomonas mating in student laboratory exercises, and for production of gamete autolysin, as well as for controls in studies of the sexual cycle. However, these strains may produce lethals in crosses and are therefore not recommended for genetic analysis.

This strain carries the nit1 and nit2 mutations and cannot grow on nitrate.

This strain is listed in some publications as NO mt+.

From Ursula Goodenough, Harvard University, June 1978

CC-620 and CC-621 are subclones of the Smith 137c wild type strain used in R.P. Levine’s laboratory at Harvard in the 1960s, selected for high mating efficiency. They are often used as demonstration material for Chlamydomonas mating in student laboratory exercises, and for production of gamete autolysin, as well as for controls in studies of the sexual cycle. However, these strains may produce lethals in crosses and are therefore not recommended for genetic analysis.

This strain carries the nit1 and nit2 mutations and cannot grow on nitrate.

From Kathleen Peterson and W.T. Ebersold, UCLA, June 1978

Phenotype: requires acetate

The ac16 mutant was isolated by Eversole, originally EV16. Krinsky and Levine found that photosynthetic electron transport measured by the Hill reaction was normal in the ac16 mutant; carotenoids were only slightly different from those of wild type cells. Levine and Goodenough listed this mutant as nonphotosynthetic, with low or absent CO2 fixation, normal pigmentation; the nature of the photosynthetic lesion is unknown.


Krinsky NI, Levine RP (1964) Carotenoids of Wild Type and Mutant Strains of the Green Aiga, Chlamydomonas reinhardi. Plant Physiol 39:680-687

Levine RP, Goodenough UW (1970) The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Annu Rev Genet 4:397-408


  • Locus:
  • AC16
  • Chromosome:
  • 10

From Kathleen Peterson and W.T. Ebersold, UCLA, June 1978

Phenotype: impaired motility

Although the motility phenotype in this strain is easy to score, for detailed analysis of pf16, CC-1034 is preferable as it has been characterized more fully. Please see that strain for more information on the PF16 locus.


  • Locus:
  • PF16
  • Chromosome:
  • 9