From Ursula Goodenough, Harvard University, March 1978

Phenotype: lacks flagella, does not mate

The bld or bald mutants were recovered as nonmating mutants in Goodenough’s laboratory, and were subsequently shown to lack flagella. The genetic lesion in this mutant has not been identified.

From Ursula Goodenough, Harvard University, March 1978

Phenotype: lacks flagella, does not mate

The bld or bald mutants were recovered as nonmating mutants in Goodenough’s laboratory, and were subsequently shown to lack flagella. The genetic lesion in this mutant has not been identified.

From Ursula Goodenough, Harvard University, March 1978

Phenotype: lacks flagella, does not mate

The bld or bald mutants were recovered as nonmating mutants in Goodenough’s laboratory, and were subsequently shown to lack flagella.

An unlinked suppressor of bld7 has been isolated by Peter Luykx. See CC-2904 through CC-2909 for Luykx’s isolates of bld7 with and without the suppressor, and the suppressor alone, all in both mating types.

Elizabeth Harris, Boynton-Gillham laboratory, Duke University

Phenotype: cold sensitive on minimal medium

This mutant is deficient in chloroplast ribosomes. Its phenotype of 70S deficiency and lack of accumulation of subunit particles resembles that of ac20. Preliminary mapping studies suggested it might be on linkage group III, but these were not followed up.

From George Witman, given to N.W. Gillham on a visit to Princeton, no accompanying letter, March 1978.

Phenotype: wall deficient

Original mutant isolated by D.R. Davies in the Ebersold/Levine 137c wild type background (equivalent to CC-125).

This is a very clean wall-deficient mutant, placed by Davies in his class C: cell walls are absent or produced in greatly reduced quantity compared to wild type. It forms flat, liquid colonies on agar, and the cells lyse in 1% solution of the nonionic detergent NP-40.

This strain was used for preparation of the DNA used for genomic sequencing.


Hyams J and Davies DR (1972) The induction and characterisation of cell wall mutants of Chlamydomonas reinhardi. Mut Res 14:381-389

Voigt J, Hinkelmann B, Harris EH (1997) Production of cell wall polypeptides by different cell wall mutants of the unicellular green alga Chlamydomonas reinhardtii. Microbiol Res 152:189-198

Boynton-Gillham laboratory, Duke University

Phenotype: antibiotic resistant (erythromycin)

This is a good isolate of the ery1b mutation, from a cross of CC-460 to a wild type strain. It is resistant to 200+ micrograms/ml erythromycin on plates. It shows only very slight growth on 50 micrograms/ml clindamycin. This strain may also contain a pf marker; very few swimming cells were observed in cultures.

This strain was used in studies by Robert Schmidt in the Boynton-Gillham laboratory in 1983-1985.

The ery1 mutations were shown by Davidson et al. (1974) to alter a protein of the large subunit of the chloroplast ribosome, which was subsequently identified by Bowers et al. as L4.


Davidson JN, Hanson MR, Bogorad L (1974) An altered chloroplast ribosomal protein in ery-M1 mutants of Chlamydomonas reinhardi. Mol Gen Genet 132:119-129

Schmidt RJ, Myers AM, Gillham NW, Boynton JE (1984) Chloroplast ribosomal proteins of Chlamydomonas synthesized in the cytoplasm are made as precursors. J Cell Biol 98:2011-2018

Bowers AK, Keller JA, Dutcher SK (2003) Molecular markers for rapidly identifying candidate genes in Chlamydomonas reinhardtii. Ery1 and ery2 encode chloroplast ribosomal proteins. Genetics 164:1345-1353


  • Locus:
  • ERY1 [PRPL4]
  • Chromosome:
  • 11

From Mike Adams in David Luck’s laboratory, Rockefeller University, April 1978

Phenotype: impaired motility

This is Luck strain pf2.13D. Strains containing the pf2 mutation swim slowly compared to wild type, but may be somewhat difficult to score.

The PF2 locus encodes component DRC4 of the dynein regulatory complex.


Rupp G and Porter ME (2003) A subunit of the dynein regulatory complex in Chlamydomonas is a homologue of a growth arrest-specific gene product. J Cell Biol 162:47-57


  • Locus:
  • PF2
  • Chromosome:
  • 11

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This mutant was isolated in Levine’s laboratory, probably by UV mutagenesis, but this is not specified in Levine’s file notes. It was found to be an allele at the AC1 locus.

When last tested, this strain was scorable by slow growth and pale green color on minimal medium. This is a reasonably good marker for linkage group VII, which is relatively sparsely populated with genetic markers.


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


  • Locus:
  • AC1
  • Chromosome:
  • 7

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac2 mutant was isolated in Levine’s laboratory. The original mutant lacked flagella and was never mapped. This strain was found to swim normally, and to be acetate-requiring and heat-sensitive.

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This mutant was isolated in Levine’s laboratory. The original mutant lacked flagella and was never mapped. This strain was found to swim normally and to have a leaky acetate requirement.

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate on low salt medium

The ac5 mutant was originally reported to form small, yellowish white colonies and to have altered thylakoid membrane stacking when grown on low salt medium. This strain is marginally scorable on Tris-minimal medium. Elevating the temperature seems to enhance the phenotype. Even so, it’s not the best genetic marker for linkage group VII; pf17 and ac1 are easier to work with.


Ebersold WT (1956) Crossing over in Chlamydomonas reinhardi. Amer J Bot 43:408-410

Goodenough UW and Staehelin LA (1971) Structural differentiation of stacked and unstacked chloroplast membranes. Freeze-etch electron microscopy of wild-type and mutant strains of Chlamydomonas. J Cell Biol 48:594-619

Levine RP and Duram HA (1973) The polypeptides of stacked and unstacked Chlamydomonas
reinhardi chloroplast membranes and their relation to photsystem II activity. Biochim Biophys Acta 325:565-572

Steinback KE and Goodenough UW (1975) Morphological and Photosynthetic Properties of Digitonin-treated Chloroplast Membranes from the Wild-type and ac-5 Strains of Chlamydomonas reinhardi. Plant Physiol. 55:864-8


  • Locus:
  • AC5
  • Chromosome:
  • 7

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac7 and ac7a mutations were isolated by Ebersold, and originally designated lg2a (see CC-673) and lg3a, respectively. They were described as acetate-requiring, and somewhat light-sensitive, having small, yellow-white, slow-growing colonies. The phenotype has become partially suppressed over the years, but at last check these strains were still scorable as yellowish green with a leaky acetate requirement, and mapped to the correct location on linkage group XI. Although it’s probably not a good choice as a mapping marker (try ery1 for this linkage group), it may be of interest as a mutant with an unknown defect in photosynthesis.age group), it may be of interest as a mutant with an unknown defect in photosynthesis.


Ebersold WT (1956) Crossing over in Chlamydomonas reinhardi. Amer J Bot 43:408-410


  • Locus:
  • AC7
  • Chromosome:
  • 11

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This is the original ac9 mutation, isolated in Levine’s laboratory.

This strain was scorable as acetate-requiring when last checked, but didn’t give terribly good results in crosses, showing incomplete tetrads and poor viability. For analysis of the non-photosynthetic phenotype (a block in psaA splicing), the allelic mutants F25 (CC-1050) and M18 (CC-1051) may be easier to work with. Of the three, M18 is the best-characterized.


Girard J, Chua NH, Bennoun P, Schmidt G, Delosme M (1980) Studies on mutants deficient in the photosystem I reaction centers in Chlamydomonas reinhardtii. Curr Genet 2:215-21


  • Locus:
  • AC9
  • Chromosome:
  • 12

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

Isolated by Eversole, originally called #140, nic-6; redefined as an acetate-requiring mutant in Levine’s laboratory based on spot tests, and listed as H8 by Ebersold et al. (1962)

This strain has been well-behaved in crosses; the acetate requirement is easy to score and it produces mostly complete tetrads. The basis for the acetate requirement hasn’t been determined, so this mutation is primarily useful as a mapping marker.


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

Girard J, Chua NH, Bennoun P, Schmidt G, Delosme M (1980) Studies on mutants deficient in the photosystem I reaction centers in Chlamydomonas reinhardtii. Curr Genet 2:215-221

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


  • Locus:
  • AC12
  • Chromosome:
  • 2

From Ursula Goodenough, Harvard University, May 1978

Isolated by Eversole, originally his #60

Like CC-522 ac12 mt+, this strain has been well-behaved in crosses, with the acetate requirement easy to score and giving mostly complete tetrads. The basis for the acetate requirement hasn’t been determined, so this is primarily useful so far as a mapping marker.


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


  • Locus:
  • AC12
  • Chromosome:
  • 2

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

At least twelve ac14 alleles were isolated in Levine’s lab at Harvard. This is the one designated simply ac14 and defines the locus. A note in the Levine files dated 5/8/63 says, ‘Robert Novak, University of Delaware, says that ac14 (+) contains cells with duplicate sets of eyespots in different locations in the cell; also location of eyespots not fixed. Also says this stock will grow on glycine as N source.’

This strain has been moderately good in past crosses, but there are better markers for this region of linkage group I, e.g. arg2, ac115, ery3.


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

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


  • Locus:
  • AC14
  • Chromosome:
  • 1

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This strain has a scorable acetate requirement, but eventually shows growth on minimal medium if the plates are left to age. There are better markers for this arm of linkage group I, e.g. arg2, ac115, ery3.

For more information on ac14, please see CC-525.


  • Locus:
  • AC14
  • Chromosome:
  • 1

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac15 mutation was UV-induced in Levine’s laboratory. The reason for the acetate requirement is unknown.

This mutant is scorable, especially at 15 degrees, but there may be some growth on minimal medium if plates are left to age.


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


  • Locus:
  • AC15
  • Chromosome:
  • 9

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This is an exceptionally clean acetate-requiring mutant, very easy to score and well-behaved in crosses. However, the reason for its stringent acetate requirement has never been determined. The original ac17 mutant was isolated in Levine’s laboratory. Many allelic mutants were also isolated, but all the strains now extant probably contain the original mutation.


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

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

Smyth RD, Martinek GW, Ebersold WT (1975) Linkage of six genes in Chlamydomonas reinhardtii and the construction of linkage test strains. J Bacteriol 124:1615-1617


  • Locus:
  • AC17
  • Chromosome:
  • 3

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate; cold sensitive on minimal medium

This strain was received as ac20 but was found by genetic analysis to be the double mutant ac20 cr1; see CC-43 for more information. It is deficient in chloroplast ribosomes and has pleiotropic defects in chloroplast structure and function. However, it does grow slightly on minimal medium. The acetate-requiring phenotype is enhanced at 15 degrees. This strain is a good choice for a strain with a general defect in chloroplast protein synthesis.


  • Locus:
  • AC20, CR1
  • Chromosome:
  • 1,12

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This strain was received as ac21, but shows linkage to the far right arm of linkage group I, suggesting that it may actually be ac24. Whatever it is, it’s a scorable marker for this region of linkage group I. However, msr1 and ery3 are easier to work with if you’re simply looking for a good marker for this region.

Levine and Goodenough listed ac24 as having normal pigments and CO2 fixation; the reason for the acetate requirement is still unknown.


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


  • Locus:
  • AC24?
  • Chromosome:
  • 1

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate, cold sensitive

The original ac30 mutant was described as pigment-deficient, having a yellow-green color. Although the color difference in CC-535 is subtle, the acetate requirement was moderately easy to score on its most recent check. Lowering the temperature to 15 degrees enhances the phenotype.


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


  • Locus:
  • AC30
  • Chromosome:
  • 6

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

ac31a was UV-induced in Levine’s laboratory. Like the allelic mutant ac31, it was originally described as forming yellow colonies.


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

Goodenough UW, Staehelin LA (1971) Structural differentiation of stacked and unstacked chloroplast membranes. Freeze-etch electron microscopy of wild-type and mutant strains of Chlamydomonas. J Cell Biol 48:594-619

Gruber HE, Rosario B (1974) Variation in eyespot ultrastructure in Chlamydomonas reinhardi (ac-31). J Cell Sci 15:481-94

Gruber HE, Rosario B (1977) Ultrastructural variations in the flagellar apparatus of Chlamydomonas reinhardii. J Submicrosc Cytol 9:103-113


  • Locus:
  • AC31
  • Chromosome:
  • 5

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This is an acetate-requiring mutant isolated in Levine’s lab, mapped to linkage group VI but otherwise poorly characterized.


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


  • Locus:
  • AC32
  • Chromosome:
  • 6

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This strain grows slightly on minimal medium and is not as easy to score on plates as the original ac40 (CC-539; please see the notes on that strain for more information on this locus).


  • Locus:
  • AC40
  • Chromosome:
  • 2

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

Both ac40 and ac40a were UV-induced in Levine’s laboratory. Of the two, ac40 is easier to score. Wollman and Bennoun showed that ac40 has an altered chlorophyll-protein complex CP 0; apparently ac40a was not tested for this property.


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

Wollman FA, Bennoun P (1982) Plant Physiol 39:889-897


  • Locus:
  • AC40
  • Chromosome:
  • 2

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This mutant is probably not the best marker for this well-populated linkage group, as the acetate requirement is difficult to score in some crosses, and many incomplete tetrads may be obtained. It may be somewhat cold-sensitive.


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:
  • AC59
  • Chromosome:
  • 12

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac80a mutation was UV-induced in Levine’s lab. This ac80 strain is difficult to score in crosses. CC-544 ac80 is better.


Givan AL, Levine RP (1967) The photosynthetic electron transport chain of Chlamydomonas reinhardi. VII. Photosynthetic phosphorylation by a mutant strain of Chlamydomonas reinhardi deficient in active P700. Plant Physiol 42:1264-1268


  • Locus:
  • AC80
  • Chromosome:
  • 2

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

The ac80 mutation was UV-induced in Levine’s lab. This mutant is deficient in P700 and photosystem I activity.

At last check, this strain showed a clean acetate requirement and was easy to score in crosses. Linkage to pf18 was confirmed.


Givan AL, Levine RP (1967) The photosynthetic electron transport chain of Chlamydomonas reinhardi. VII. Photosynthetic phosphorylation by a mutant strain of Chlamydomonas reinhardi deficient in active P700. Plant Physiol 42:1264-8

Givan AL, Levine RP (1969) The photosynthetic electron transport chain of a mutant strain of Chlamydomonas reinhardi lacking P700 activity. Biochim Biophys Acta 189:404-10

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


  • Locus:
  • AC80
  • Chromosome:
  • 2

From Ursula Goodenough, Harvard University, May 1978

Phenotype: requires acetate

This is an allele at the AC115 locus, UV-induced in Levine’s laboratory. Please see CC-548 for more information on ac115.


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