Born:  March 16, 1943, New York City

 Five children (b. 1970, 1974, 1980, 1982, 1985)


Current Position:

     Professor of Biology Emerita

     Department of Biology

     Washington University

     St. Louis, MO 63130



     Radcliffe College, 1960-1961, Ann Radcliffe Scholar.

     Barnard College, 1961-1963, A.B. cum laude with honors in Zoology,

          Phi Beta Kappa, von Wahl Prize in Zoology

     Columbia University, 1963-1965, M.A. in Zoology

          (Advisor:  Professor Lee D. Peachey)

     Harvard University, 1965-1969, Ph.D. in Biology

          (Thesis Advisor:  Professor Keith R. Porter)


Positions Held:

     1982-2017  Professor of Biology, Washington University

     1978-1982  Associate Professor of Biology, Washington University

     1976-1978  Associate Professor of Biology, Harvard University

     1971-1976  Assistant Professor of Biology, Harvard University

     1969-1971  NIH Post-Doctoral Fellow, Harvard University

                  (sponsor:  Professor Lawrence Bogorad)

     1964-1969  NIH Pre-Doctoral Fellow, Columbia and Harvard Universities

     1963-1964  Eugene Higgins Fellow, Columbia University


Professional Activities:

     1978-1982 Member, Molecular Cytology Study Section, NIH

     1987-1992 Member, Cell Biology Review Panel, NSF

     1979-1983 Associate Editor, Journal of Cell Biology

     1994-1998 Associate Editor, Cell Motility and the Cytoskeleton

     1997-2000 Associate Editor, Journal of Phycology     

     1988      Coordinator, Meeting on Cell and Molecular Biology of 

                  Chlamydomonas, Cold Spring Harbor Laboratory 

     1990      Coordinator, Regional Meeting of the American Society for 

                  Cell Biology, Chicago

     1990-1993 Council Member, American Society for Cell Biology (ASCB)

     1990-1993 Member, Education Committee, ASCB

     1992-1993 Chair, Women in Cell Biology Committee, ASCB

     1994-1995 President, ASCB

     1994-1995 Member, Future of Space Science Steering Group (FOSS), Space 

                  Studies Board, National Research Council (NRC)     

     1995-1998 Member, Commission on Life Sciences, NRC

     1996-1998 Adjunct Member, Trends in Early Research Careers, Biology 

                  Board, NRC

     1996-2010 President, Phi Beta Kappa, Washington University chapter

     1999-2000 Phi Beta Kappa Visiting Scholar

     1999-2002 Board member, Science Writers Fellowship Program, Marine 

                  Biological Laboratory 

     2001-2003 Editor, Eukaryotic Cell

     2002 Coordinator, Meeting on Cell and Molecular Biology of  

                   Chlamydomonas, Vancouver, Canada

     2003-2009 Chair, Women in Cell Biology Committee, ASCB

     2012 –    Editorial Board, Algal Research

     2012-2018  Scientific Advisory Committee, Integrated Microbial Diversity,

    Canadian Institute for Advanced Research      

Science Policy:

     1992-2009 Member, Public Policy Committee, ASCB

     1995-1997 Member, Joint Steering Committee for Public Policy

     1993-1995 Chair, NSF and NASA Committees, FASEB Consensus Conferences

                  on Public Policy

     1997-1999 Member, FASEB Subcommittee on Ethical Issues of Genetic Research


Washington University Teaching

            Biology 334, Introduction to Cell Biology (junior-level) 

            Biology/Earth and Planetary Sciences/Physics 210, Epic of Evolution (non-science majors)

            Biology 4331: Algae: Cell Biology and Molecular Evolution (graduate 


            Biology 5702: Current Approaches in Plant Research (course coordinator)



Faculty Teaching Awards, Washington University, 1986 and 1994

Founder’s Day Distinguished Faculty Award, Washington University, 1995

Senior Career Recognition Award, American Society for Cell Biology, 1999

Fellow, American Women in Science, 2009

Fellow, American Academy of Arts and Sciences, 2009

Fellow, American Academy of Microbiology, 2013

Doctor of Letters Honoris Causa, Meadville School of Theology, 2022

Faculty Teaching Awards, Washington University, 1986 and 1994

Founder’s Day Distinguished Faculty Award, Washington University, 1995

Senior Career Recognition Award, American Society for Cell Biology, 1999

Fellow, American Women in Science, 2009

Fellow, American Academy of Arts and Sciences, 2009

Fellow, American Academy of Microbiology, 2013

Doctor of Letters Honoris Causa, Meadville School of Theology, 2022

Member, National Academy of Sciences, 2023


Research Interests and Support:

Research has focused on the life cycle of the green alga Chlamydomonas reinhardtii using molecular-genetic and ultrastructural analysis. Additional projects have analyzed the ciliary axoneme, the cell walls of various algae, the formation of algal lipid bodies, and the ultrastructure of lichens. The lab has been continuously supported by grants from the NIH, NSF, USDA, and DOE.


Research Articles:

Johnson (former surname), U.G., and K.R. Porter. 1968. Fine structure of cell division in Chlamydomonas reinhardi.  Basal bodies and microtubules. J. Cell Biol. 38:403-426.

Goodenough, U.W. and R.P. Levine. 1969. Chloroplast ultrastructure in mutant strains of Chlamydomonas reinhardi lacking components of the photosynthetic apparatus. Plant Physiol. 44:990-1000.

Goodenough, U.W., J.J. Armstrong, and R.P. Levine. 1969. Photosynthetic properties of ac-31, a mutant strain of Chlamydomonas reinhardi devoid of chloroplast membrane stacking. Plant Physiol. 44:1001-1012.

Goodenough, U.W. and R.P. Levine. 1970. Chloroplast structure and function in ac-20, a mutant strain of 

             Chlamydomonas reinhardi.  III.  Chloroplast ribosomes and membrane organization. J. Cell Biol.


Goodenough, U.W. 1970. Chloroplast division and pyrenoid formation in Chlamydomonas reinhardi. J. Phycol. 6:1-6.

Goodenough, U.W. and L.A. Staehelin. 1971. Structural differentiation of stacked and unstacked chloroplast membranes:  Freeze-etch electron microscopy of wild type and mutant strains of Chlamydomonas reinhardi. J. Cell Biol. 48:594-619.

Goodenough, U.W. 1971. The effects of inhibitors of RNA and protein synthesis on chloroplast structure and function in Chlamydomonas reinhardi. J. Cell Biol. 50:35-49.

Goodenough, U.W. and R.P. Levine. 1971. The effects of inhibitors of RNA and protein synthesis on the recovery of chloroplast ribosomes, membrane organization and photosynthetic electron transport in the ac-20 strain of Chlamydomonas reinhardi. J. Cell Biol. 50:50-62.

Steinback, K.E. and U.W. Goodenough. 1975. Morphological and photosynthetic properties of digitonin-treated chloroplast membranes from the wild-type and ac-5 strains of Chlamydomonas reinhardi.  Plant Physiol. 55:863-869.

Martin, N.C. and U.W. Goodenough. 1975. Gametic differentiation in Chlamydomonas reinhardi.  I.  Production of gametes and their fine structure. J. Cell Biol. 67:587-605.

Bergman, K., U.W. Goodenough, D.A. Goodenough, J. Jawitz, and H. Martin. 1975. Gametic differentiation in Chlamydomonas reinhardi.  II.  Flagellar membranes and the agglutination reaction. J. Cell Biol. 67:606-622.

Goodenough, U.W. and R.L. Weiss. 1975. Gametic differentiation in Chlamydomonas reinhardi.  III.  Cell wall lysis and microfilament-associated mating structure activation in wild-type and mutant strains. J. Cell Biol. 67:623-637.

Goodenough, U.W. and H.S. St. Clair. 1975. Bald-2:  A mutation affecting the formation of doublet and triplet 

             sets of microtubules in Chlamydomonas reinhardi. J. Cell Biol. 66:480-491.

Goodenough, U.W., C. Hwang, and H. Martin. 1976. Isolation and genetic analysis of mutant strains of Chlamydomonas reinhardi defective in gametic differentiation. Genetics 82:169-186.

Martin, N.C., K.S. Chiang, and U.W. Goodenough. 1976. Turnover of chloroplast and cytoplasmic ribosomes during gametogenesis in Chlamydomonas reinhardi. Dev. Biol. 51:190-201.

Weiss, R.L., D.A. Goodenough, and U.W. Goodenough. 1977. Membrane particle arrays associated with the basal body and with contractile-vacuole secretion in Chlamydomonas. J. Cell Biol. 72:133-143.

Weiss, R.L., D.A. Goodenough, and U.W. Goodenough. 1977. Membrane differentiations at sites specialized for cell fusion. J. Cell Biol. 72:144-160.

Goodenough, U.W. and R.L. Weiss. 1978. Interrelationships between microtubules, a striated fiber, and the gametic mating structure of Chlamydomonas reinhardi.  J. Cell Biol. 76:430-438.

Minami, S.A. and U.W. Goodenough. 1978. Novel polypeptide synthesis induced by gametic cell fusion in Chlamydomonas reinhardi.  J. Cell Biol. 77:165-181.

Goodenough, U.W., Hwang, C., and A.J. Warren.  1978.  Sex-limited expression of gene loci controlling flagellar membrane agglutination in the Chlamydomonas mating reaction.  Genetics 89:235-243.

Forest, C., D.A. Goodenough, and U.W. Goodenough. 1978. Flagellar membrane agglutination and sexual signaling in the conditional gam-1 mutant of Chlamydomonas. J. Cell Biol. 79:74-84.

Goodenough, U.W. and D. Jurivich. 1978. Tipping and mating-structure activation induced in Chlamydomonas gametes by flagellar-membrane antisera. J. Cell Biol. 79:680-693.

Adair, W.S., D. Jurivich, and U.W. Goodenough. 1978. Localization of cellular antigens in sodium dodecyl sulfate-polyacrylamide gels. J. Cell Biol. 79:281-285.

Mesland, D.A.M., J.L. Hoffman, E. Caligor and U.W. Goodenough. 1980. Flagellar tip activation stimulated by membrane adhesions in Chlamydomonas gametes. J. Cell Biol. 84:599-617.

Hoffman, J.L. and U.W. Goodenough. 1980. Experimental dissection of flagellar surface motility in Chlamydomonas.  J. Cell Biol. 86:656-665.

Hwang, C., B.C. Monk, and U.W. Goodenough. 1981. Linkage of mutations affecting minus flagellar membrane agglutinability to the mtmating type locus of Chlamydomonas. Genetics 99:41-47.

Goodenough, U.W., P.A. Detmers, and C. Hwang. 1982. Activation for cell fusion in Chlamydomonas:  Analysis of wild-type gametes and non-fusing mutants. J. Cell Biol. 42:378-386.

Adair, W.S., B.C. Monk, R. Cohen, C. Hwang, and U.W. Goodenough. 1982. Sexual agglutinins from the Chlamydomonas flagellar membrane:  Partial purification and characterization. J. Biol. Chem. 257:4593-4602.

Adair, W.S.* 1982. The spaghetti overlay:  Simultaneous screening of multiple polyclonal and monoclonal antibodies by immunoautoradiography.  Anal. Biochem. 125:299-306.

Goodenough, U.W., and J.E. Heuser. 1982. Substructure of the outer dynein arm. J. Cell Biol. 95:798-815. 

Goodenough, U.W. 1983. Motile detergent-extracted cells of Tetrahymena and Chlamydomonas. J. Cell Biol. 96:1610-1621.

Adair, W.S., C. Hwang, and U.W. Goodenough. 1983. Identification and visualization of the sexual agglutinin from mating-type plus flagellar membranes of Chlamydomonas. Cell 33:183-193.

Hourcade, D.* 1983. Marker rescue from bleomycin-treated Chlamydomonas reinhardi.  Genetics 104:391-404.

Detmers, P., U.W. Goodenough, and J. Condeelis.  1983.  Elongation of the fertilization tubule in Chlamydomonas:  New observations on the core micro-filaments and the effects of transient intracellular signals on their structural integrity.  J. Cell Biol. 97:522-532.

Monk, B.C., W.S. Adair, R.A. Cohen, and U.W. Goodenough. 1983. Topography of Chlamydomonas:  Fine structure and polypeptide components of the gametic flagellar membrane surface and the cell wall.  Planta 158:517-533.

Cooper, J.B., W.S. Adair, R.P. Mecham, J.E. Heuser, and U.W. Goodenough. 1983. The Chlamydomonas agglutinin is a hydroxyproline-rich glycoprotein. Proc. Natl. Acad. Sci. USA 80:5898-5901.

Collin-Osdoby, P., W.S. Adair, and U.W. Goodenough. 1984. Chlamydomonas agglutinin conjugated to agarose beads as an in vitro probe of adhesion. Exp. Cell Res. 150:282-291.

Goodenough, U.W., and J.E. Heuser. 1984. Structural comparison of purified dynein proteins with in situ dynein arms. J. Mol. Biol. 180:1083-1118.

Galloway, R.E., and L.R. Holden.* 1984. Transmission and recombination of chloroplast genes in asexual crosses of Chlamydomonas reinhardii I. Flagellar agglutination prior to fusion does not promote uniparental inheritance or affect recombinant frequencies. Curr. Genet. 8:399-405.

* Post-doctoral fellows reporting independently executed project in Goodenough lab.

Goodenough, U.W., and J.E. Heuser. 1985. Substructure of inner dynein arms, radial spokes, and the central pair/projection complex of cilia and flagella. J. Cell Biol. 100:2008-2018.

Goodenough, U.W., W.S. Adair, P. Collin-Osdoby, and J.E. Heuser. 1985. Structure of Chlamydomonas agglutinin and related flagellar surface proteins in situ and in vitro.  J. Cell Biol. 101:924-942.

Collin-Osdoby, P., and W.S. Adair.* 1985. Characterization of the purified Chlamydomonas minus agglutinin.  J. Cell Biol. 101:1144-1152.   

Sale, W.S., U.W. Goodenough, and J.E. Heuser. 1985. The substructure of isolated and in situ outer dynein arms of sea urchin sperm flagella. J. Cell Biol. 101:1400-1412.

Galloway, R.E., and U.W. Goodenough. 1985. Genetic analysis of mating locus-linked mutations in Chlamydomonas reinhardi. Genetics 111:447-461.

Goodenough, U.W., and J.E. Heuser. 1985. The Chlamydomonas cell wall and its constituent proteins analyzed by the quick-freeze deep-etch technique. J. Cell Biol. 101:1550-1568.

Goodenough, U.W. 1986. Experimental analysis of the adhesion reaction between isolated Chlamydomonas flagella.  Exp. Cell. Res. 166:237-246.

Goodenough, U.W., B. Gebhart, R. Mecham, and J.E. Heuser. 1986. Crystals of the Chlamydomonas 

 reinhardi cell wall:  Polymerization, depolymerization, and purification of glycoprotein monomers. J. 

Cell. Biol. 103:405-417.

Goodenough, U.W., B. Gebhart, V. Mermall, D. Mitchell, and J.E. Heuser. 1987. HPLC fractionation of Chlamydomonas dynein extracts and characterization of three inner-arm dyneins. J. Mol. Biol. 194:481-494.

Ferris, P.J., and U.W. Goodenough. 1987. Transcription of novel genes, including a gene linked to mating-type, induced by Chlamydomonas fertilization. Molec. Cell. Biol. 7:2360-2366.

Pasquale, S.M., and U.W. Goodenough. 1987. Cyclic AMP functions as a primary sexual signal in gametes of Chlamydomonas reinhardtii. J. Cell Biol. 105:2279-2292.

Adair, W.S., S.A. Steinmetz, D.M. Mattson, U.W. Goodenough, and J.E. Heuser. 1987. Nucleated assembly of Chlamydomonas and Volvox cell walls. J. Cell Biol. 105:2373-2382.

Goodenough, U.W., and J.E. Heuser. 1988. Molecular organization of cell-wall crystals from Chlamydomonas reinhardtii and Volvox carteri. J. Cell Sci. 90:717-733.

Goodenough, U.W., and J.E. Heuser. 1988. Molecular organization of the cell wall and cell-wall crystals from Chlamydomonas eugametos. J. Cell Sci. 90:735-750.

Goodenough, U.W., and J.E. Heuser. 1988. Structure of soluble and in situ ciliary dyneins visualized by quick-freeze deep-etch microscopy.  In: Cell Movement, Vol. I. The Dynein ATPases. F.D. Warner, P. Satir, and I.R. Gibbons, eds. New York, Alan Liss. pp. 121-140.

Ferris, P.J.* 1989. Characterization of a Chlamydomonas transposon, Gulliver, resembling those in higher plants. Genetics 122:363-377.

Goodenough, U.W. 1989. Cyclic AMP enhances the sexual agglutinability of Chlamydomonas flagella. J. Cell Biol. 109:247-252.

Woessner, J.P., and U.W. Goodenough. 1989. Molecular characterization of a zygote wall protein: an extensin-like molecule in Chlamydomonas reinhardtii. The Plant Cell 1:901-911.

Woessner, J.P., and U.W. Goodenough. 1992. Zygote and vegetative cell wall proteins in Chlamydomonas reinhardtii share a common epitope, (Ser Pro)x. Plant Sciences 83:65-76.

Matters, G.L., and U.W. Goodenough. 1992. A gene/pseudogene tandem duplication encodes a cysteine-rich protein expressed during zygote development in Chlamydomonas reinhardtii. Mol. Gen. Genet. 232:81-88.

Goodenough, U.W. 1993. Tipping of flagellar agglutinins by gametes of Chlamydomonas reinhardtii. Cell Motil. Cytoskel. 25:179-189.

Goodenough, U.W., B. Shames, L. Small, T. Saito, R. C. Crain, M.A. Sanders and J.L. Salisbury. 1993. The role of calcium in the Chlamydomonas reinhardtii mating reaction. J. Cell Biol. 121:365-374.

Saito, T., L. Small, and U.W. Goodenough. 1993. Activation of adenylyl cyclase in Chlamydomonas reinhardtii by adhesion and by heat. J. Cell Biol. 122:137-147.

Waffenschmidt, S., J.P. Woessner, K. Beer, and U.W. Goodenough. 1993.  Evidence that isodityrosine crosslinking mediates the insolubilization of cell-wall HRGPs in Chlamydomonas. The Plant Cell 5:809-820.

Armbrust, E.V., P.J. Ferris, and U.W. Goodenough. 1993. A mating type-linked gene cluster expressed in Chlamydomonas participates in the uniparental inheritance of the chloroplast genome.  Cell 74:801-811.  

Ferris, P.J., and U.W. Goodenough. 1994. The mating-type locus of Chlamydomonas reinhardtii contains highly rearranged DNA sequences. Cell 76:1135-1145.

Woessner, J.P., A.J. Molendijk, P. van Egmond, F.M. Klis, U.W. Goodenough, and M.A. Haring. 1994. Domain conservation in several volvocalean cell wall proteins. Plant Molec. Biol. 26:947-60.

Campbell, A.M., H.J. Rayala, and U.W. Goodenough. 1995. The isol gene of Chlamydomonas is involved in sex determination. Mol. Biol. Cell 6:87-95.

Ferris, P.J.* 1995. Localization of the nic-7, ac-29, and thi-10 genes within the mating-type locus of Chlamydomonas reinhardtii.  Genetics 141:543-549.

Armbrust, E.V., A. Ibrahim, and U.W. Goodenough. 1995. A mating type-linked mutation that disrupts the uniparental inheritance of chloroplast DNA also disrupts cell-size control in Chlamydomonas. Mol. Biol. Cell 6:1807-1818.

Ferris, P.J., J.P. Woessner, and U.W. Goodenough. 1996. A sex recognition glycoprotein is encoded by the plus mating-type gene fus1 of Chlamydomonas reinhardtii. Mol. Biol. Cell 7:1235-1248.

Ferris, P.J., and U.W. Goodenough. 1997.  Mating type in Chlamydomonas is specified by mid, the minus-dominance gene. Genetics 146::859-869.

Ferris, P.J., C. Pavlovic, S. Fabry, and U.W. Goodenough. 1997. Rapid evolution of sex-related genes in Chlamydomonas.  Proc. Natl. Acad. Sci. USA 94:8634-8639.

Goodenough, U.W., and J.E. Heuser. 1999. Deep-etch analysis of adhesion complexes between gametic flagellar membranes of Chlamydomonas reinhardtii (Chlorophyceae). J. Phycol. 35:756-767.

Suzuki, L., J.P. Woessner, H. Uchida, H. Kuroiwa, Y. Yuasa, S. Waffenschmidt, U. Goodenough, and T. Kuroiwa. 2000. A zygote-specific protein with hydroxyproline-rich glycoprotin domains and lectin-like domains involved in the assembly of the cell wall of Chlamydomonas reinhardtii (Chlorophyta). J. Phycol. 36:571-583.

Ferris, P.J., J.P. Woessner, S. Waffenschmidt, S. Kilz, J. Drees, and U.W. Goodenough. 2001. Glycosylated polyproline II rods with kinks as a structural motif in plant hydroxyproline-rich glycoproteins.  Biochemistry 40:2978-2987.

Umen, J.G., and U.W. Goodenough. 2001. Control of cell division by a retinoblastoma protein homolog in Chlamydomonas. Genes & Development 15:1652-1661.

Umen, J.G., and U. W. Goodenough. 2001. Chloroplast DNA methylation and inheritance in Chlamydomonas.  Genes & Development 15:2585-2597.  

Ferris, P.J., E.V. Armbrust, and U.W. Goodenough.  2002.  Genetic structure of the mating-type locus of Chlamydomonas reinhardtii.  Genetics 160:181-200.

Ferris, P.J., S. Waffenschmidt, J.G. Umen, H. Lin, J-H. Lee, K. Ishida, T. Kubo, J. Lau, and U.W. Goodenough. 2005. Plus and minus sexual agglutinins from Chlamydomonas reinhardtii. Plant Cell 17:597-615.

Lin, H. and U.W. Goodenough. 2007. Gametogenesis in the Chlamydomonas reinhardtii minus mating type is controlled by two genes, MID and MTD1. Genetics 176:913-925.

Lee, J-H., S. Waffenschmidt, L. Small, and U.W. Goodenough. 2007. Between-species analysis of short-repeat modules in cell wall and sex-related hydroxyproline-rich glycoproteins of Chlamydomonas. Plant Physiol. 144:1813-1826.

Lee, J-H., H. Lin, S. Joo, and U. Goodenough. 2008. Early sexual origins of homeoprotein heterodimerization and evolution of the plant KNOX/BELL family. Cell 133:829-840.

Worden, A.Z., J-H. Lee …(46 authors)…U.W. Goodenough, Y. Van de Peer, and I.V. Grigoriev. 2009. Green evolution and dynamic adaptations revealed by genomes of the marine picoeukaryotes Micromonas.  Science 324:268-272.

Wang, Z.T., N. Ullrich, S. Joo, S. Waffenschmidt, and U. Goodenough. 2009. Algal lipid bodies: Stress induction, purification, and biochemical characterization in wild-type and starch-less Chlamydomonas reinhardtii. Eukaryotic Cell 8:1856-1868.

Goodson, C., R. Roth, Z.T. Wang, and U. Goodenough. 2011. Structural correlates of cytoplasmic and chloroplast lipid body synthesis in Chlamydomonas reinhardtii and stimulation of lipid body production with acetate boost. Eukaryotic Cell 10:1592-1606.

Weiss, T.L., R. Roth, C. Goodson, S. Vitha, I. Black, P. Azadi, J. Rusch, A.Holzenburg, T. Devarenne, and U.

  Goodenough. 2012. Colony organization in the green alga Botryococcus braunii (Race B) is specified

by a complex extracellular matrix. Eukaryotic Cell 11:1424-1440.

Goodenough, U., I. Blaby, D. Casero, S.D. Gallaher, C. Goodson, S. Johnson, J-H. Lee, S. S. Merchant, M.

Pellegrini, R. Roth, J. Rusch, M. Singh, J. G. Umen, T. L. Weiss, and T. Wulan. 2014. The path to

triacylglyceride obesity in the sta6 strain of Chlamydomonas reinhardtii. Eukaryotic Cell


Ramundo, S., D. Casero, T. Mühlhaus, D. Hemme, F. Sommer, M. Crèvecoeur, M.Rahire, M. Schroda, J. 

Rusch, U. Goodenough, M. Pellegrini, M. E. Perez-Perez, J. L. Crespo, O. Schaad, N. Civic and J. D. 

Rochaix. 2014. Conditional depletion of the chloroplast ClpP1 protein activates nuclear genes involved in autophagy and plastid protein quality control. Plant Cell 26:2201-2222.

Allen, A., C. Dupont, R. Valas, J. McCrow, J. Moustafa, A. Walworth, U. Goodenough, R. Roth, S. Hogle, J. 

Bai, Z. Johnson, E. Mann, B. Palenik, K. Barbeau, and J.C. Venter. 2014. Genomes and gene

 expression across light and productivity gradients in eastern subtropical Pacific microbial communities. ISME Journal 9:1076-1092.

Scholz, M.J., T.L. Weiss, R.E. Jinkerson, J. Jing, R. Roth, U. Goodenough, M.C. Posewitz, and H.G. Gerken.

2014. Ultrastructure and composition of the Nannochloropsis gaditana cell wall. Eukaryotic Cell 13: 1450-1464.

Lee, J-H., J.E. Heuser, R. Roth, and U. Goodenough. 2015. Eisosome ultrastructure and evolution in fungi, microalgae, and lichens. Eukaryotic Cell 14:1017-1042.

Mackinder, L.C.M., M.T. Meyer, T. Mettler-Altmann, V. Chen, M.C. Mitchell, O. Caspari, E.S.F. Rosenzweig,

L. Pallesen, G.Reever, A. Itakura,R. Roth, F. Sommer, S.Geimer, T. Mühlhaus, M. Schroda, U. Goodenough, M. Stitt, H. Griffiths, and M.C. Jonikas. 2016. Repeat protein links Rubisco to form the eukaryotic carbon-concentrating organelle. Proc. Natl.Acad. Sci. USA 113: 5958-5963.

Dinc, E., L. Tian, L.M. Roy, R. Roth, U. Goodenough, and R. Croce. 2016. LHCSR1 induces a fast and  reversible pH-dependent fluorescence quenching in LHCII in Chlamydomonas reinhardtii cells. Proc. Natl. Acad. Sci. USA 113: 7673-7678.

McCarthy, J., S. Smith, J. McCrow, M. Tan, H. Zheng, K. Beeri, R. Roth, C. Lichtle, U. Goodenough, C. Bowler, C. Dupont, and A. Allen. 2017. Nitrate reductase knockout uncouples nitrate transport from nitrate assimilation and drives repartitioning of carbon flux in a model pennate diatom. The Plant Cell  29: 2047-2070.

Limardo, A.J., S. Sudek, J.C. Chang, C. Poirier, Y.M Rii, M.Blum, R. Roth, U. Goodenough, M.J. Church, and

A.Z. Worden. 2017. Quantitative biogeography of picoprasinophytes establishes ecotype distributions and significant contributions to marine phytoplankton. Environmental Microbiol. 19: 3219- 3234.

Wolf, B.M., D.M. Niedzwiedzki, N.C.M. Magdaong, R. Roth, U. Goodenough, and R.E. Blankenship. 2018. Characterization of a newly isolated freshwater Eustigmatophyte alga capable of utilizing far-red light as its sole light source. Photosynth. Res. 135: 177-187.

Guo, J., S. Wilken, V. Jimenez, C.J. Choi,, C. Ansong, R. Dannebaum, L.Sudek, D.S. Milner, C. Bachy, E.N.

Reistetter, V.A. Elrod, D. Klimov, S.O. Purvine, C.L. Wei, G. Kunde-Ramamoorthy, T.A. Richards, U. Goodenough, R.D. Smith, S.J. Callister, and A.Z. Worden. 2018. Specialized proteomic responses and an ancient photoprotection mechanism sustain marine green algal growth during phosphate limitation. Nature Microbiol. 3:781-790.

Goodenough, U., R. Roth, T. Kariyawasam, A. He, and J-H Lee. 2018. Epiplasts: Membrane skeletons and 

epiplastin proteins in euglenids, glaucophytes, cryptophytes, ciliates, dinoflagellates, and apicomplexans. mBio 9: e02020.

Price, D.C., U. Goodenough, R. Roth, J-H. Lee, F. Facchinelli, S.G. Ball, U. Cenci, C.X. Chan, N.E. Wagner,

H.S. Yoon, A.P.M Weber, and D. Bhattacharya. 2019. Analysis of an improved Cyanophora

paradoxa genome assembly. DNA Res. 26: 287-299

Goodenough, U., A. A. Heiss, R. Roth, J. Rusch, and J-H. Lee. 2019. Acidocalcisomes: Ultrastructure, biogenesis, and distribution in microbial eukaryotes. Protist 170: 287-313.

Kariyawasam, T., S. Joo, U. Goodenough, and J-H. Lee. 2019. Novel approaches for generating and manipulating diploid strains of Chlamydomonas reinhardtii. Algae 34:35-43.

Ikatura, A., K.X. Chan, N. Atkinson, L. Pallesen, L. Wang, G. Reeves, W. Patena, O. Caspari, R. Roth, U. Goodenough, A. McCormick, H. Griffiths, and M. Jonikas. 2019. A Rubisco-binding protein is required for normal pyrenoid number, tubule content, and starch sheath shape in Chlamydomonas reinhardtii. Proc. Natl. Acad. Sci. USA 37:18445-18454.

Polle, J.E.W., R. Roth, A. Ben-Amotz, and U. Goodenough. 2020. Ultrastructure of the green alga Dunaliella salina strain CCAP19/18 (Chlorophyta) as investigated by quick-freeze deep-etch electron microscopy. Algal Res. 49: 101953.

Goodenough, U. 2021. Introduction to the lichen ultrastructure series. Algal Res. 59: 102460.

Roth, R., and U. Goodenough. 2021. Lichen 1. Solo fungal and algal partners. Algal Res. 58: 102334.

Goodenough, U., and R. Roth. 2021. Lichen 2. Constituents. Algal Res.58: 102356.

Roth, R., R. Wagner, and U. Goodenough. 2021. Lichen 3. Outer layers. Algal Res. 60: 102462.

Goodenough, U., R. Wagner, and R. Roth. 2021. Lichen 4. The algal layer. Algal Res. 58: 102355.

Goodenough, U., and R. Roth. 2021. Lichen 5. Medullary and biofilm layers. Algal Res. 58: 102333.

Arakawa, S., T. Kanaseki, R. Wagner, and U. Goodenough. 2022. Ultrastructure of the foliose lichen

Myelochroa leucotiliza and its solo fungal and algal (Trebouxia sp.) partners. Algal Res. 62:102571.

Joo, S., T. Kariyawasam, M. Kim, E. Jin, U. Goodenough, and J-H. Lee. 2022. Sex-linked deubiquitinase

establishes uniparental transmission of chloroplast DNA. Nature Commun. 13: 1133. 


Review Articles, Chapters and Books:


Surzycki, S.J., U.W. Goodenough, R.P. Levine and J.J. Armstrong. 1970. Nuclear and chloroplast control of chloroplast structure and function in Chlamydomonas reinhardi.  Symp. Soc. Exp. Biol. 24:13-35.

Goodenough, U.W., R.K. Togaski, A. Paszewski and R.P. Levine. 1970. Inhibition of chloroplast ribosome formation by gene mutation in Chlamydomonas reinhardi.  Symposium on the Biogenesis of Mitochondria and Chloroplasts, Canberra, Australia, pp. 224-234.

Levine, R.P. and U.W. Goodenough. 1970. The genetics of photosynthesis and of the chloroplast in Chlamydomonas reinhardi. Ann. Rev. Genetics 49:397-408. 

Goodenough, U.W. and R.P. Levine. 1970. The genetic activity of mitochondria and chloroplasts.  Scientific American 233:22-29.

Goodenough, U.W., and R.P. Levine. 1972. Genetics. New York, Holt, Reinhardt, and Winston.

Goodenough, U.W. and R.P. Levine.  1974.  Methods for the study of chloroplast membranes of wild-type and mutant strains of Chlamydomonas reinhardi.  In: Methods in Enzymology, XXXII, S. Fleischer and L. Packer, eds., Academic Press. pp. 871-880.

Goodenough, U.W. 1976. Genetics, 2d edition.  New York, Holt/Saunders. 840 pp

Goodenough, U.W. 1977. Mating interactions in Chlamydomonas.  In:  Microbial Interactions (Receptors and Recognition, Series B, Volume 3), J.L. Reissig, ed.  Chapman and Hall, Ltd.  pp. 325-350.

Goodenough, U.W. and C. Forest. 1978. Genetics of cell-cell interactions in Chlamydomonas.  In:  The Molecular Basis of Cell-Cell Interaction.  R.A. Lerner and D. Bergsma, eds.  New York, Alan Liss. pp. 429-438.

Goodenough, U.W., W.S. Adair, E. Caligor, C.L. Forest, J.L. Hoffman, D.A.M. Mesland, and S. Spath.  1980.  Membrane-membrane and membrane-ligand interactions during Chlamydomonas mating.  In:  Membrane-Membrane Interactions.  N.B. Gilula, ed.  New York.  Raven Press. pp. 131-152.

Goodenough, U.W.  1980.  Sexual Microbiology:  The mating reactions of Chlamydomonas, Tetrahymena and Saccharomyces and their genetic control. Symp. Soc. Gen. Microbiol. 30:301-328.

Goodenough, U.W. and S. Adair.  1980.  Membrane adhesions between Chlamydomonas Mediator of Developmental Processes.  S. Subtelny and N.K. Wessells, eds. N.Y. Academic Press. pp. 101-112.

Goodenough, U.W., and J. Thorner.  1983.  Sexual differentiation and mating strategies in the yeast Saccharomyces and in the green alga Chlamydomonas. In:  Cell Interactions and Development:  Molecular Mechanisms.  K.M. Yamada, ed.  New York, John Wiley. pp. 29-75.

Goodenough, U.W.  1984.  Genetics, 3rd edition.  New York, Holt/Saunders. 930 pp.

Goodenough, U.W.  1985.  Origins and evolution of eukaryotic sex.  In: Origins and Evolution of Sex.  H.O. Halvorson and A. Monroy, eds.  N.Y., Alan R. Liss, Inc. pp. 123-140.

Goodenough, U.W., W.S. Adair, P. Collin-Osdoby, and J.E. Heuser.  1985. Chlamydomonas cells in contact.  In:  The Cell in Contact.  G.M. Edelman and J.P. Thiery, eds.  N.Y., John Wiley. pp. 111-135.

Adair, W.S.* 1985. Characterization of Chlamydomonas sexual agglutinins. J. Cell Sci. Suppl. 2:233-260.

Goodenough, U.W., and J.E. Heuser. 1985. Outer and inner arms of cilia and flagella.  Cell 41:341-342.

Goodenough, U.W., and P.J. Ferris. 1987. Genetic regulation of development in Chlamydomonas. In: Genetic Regulation of Development. W.L. Loomis, ed. N.Y., Alan Liss Inc. pp. 171-190.

Goodenough, U.W. and W.S. Adair. 1989. Recognition proteins of Chlamydomonas reinhardtii (Chlorophyceae).  In: Algae as Experimental Systems.  A.W. Coleman, L.J. Goff, and J.R. Stein-Taylor, eds. New York, Alan R. Liss, Inc. pp. 171-185.

Goodenough, U.W. 1989. Cilia, flagella, and the basal apparatus.  Curr. Opinion Cell Biol. 1:58-62.

Goodenough, U.W. 1989. Basal body chromosomes? Cell 59:1-3.

Goodenough, U.W. 1991. Deception by pathogens. Amer. Sci. 79:344-356.

Goodenough, U.W., 1991.  Chlamydomonas mating interactions.  In: Microbial Cell-Cell Interactions. M. Dworkin, ed., Washington, D.C. American Society for Microbiology. pp. 71-112.

Goodenough, U.W. 1992. Green yeast. Cell 70:533-538.

Goodenough, U.W. and T. Platt, eds. 1992. Prokaryotes and lower eukaryotes. Curr. Opinion Genet. Devel. 2(5).

Woessner, J.P., and U.W. Goodenough. 1994.  Volvocine cell walls and their constituent glycoproteins: An evolutionary perspective. Protoplasma 181:245-258.

Goodenough, U.W., E.V. Armbrust, A.M. Campbell, and P.J. Ferris. 1995. Molecular genetics of sexuality in Chlamydomonas. Ann. Rev. Plant Physiol. Plant Mol. Biol. 46:21-44.

Goodenough, U., H. Lin, and J-H Lee. 2007. Sex determination in Chlamydomonas. Sem. Cell Devel. Biol. 18: 350-361.

Snell, W.J. and U.W. Goodenough. 2008. Flagellar adhesion, flagellar-generated signaling, and gamete fusion during mating.  In: Chlamydomonas Sourcebook, volume 3, Flagella.  G. Witman, ed. Amsterdam: Elsevier. pp. 369-394. 

Goodenough, U. and J. Heitman. 2014. Origins of eukaryotic sexual reproduction. In: Origin and Evolution of Eukaryotes. P. Keeling and E. Koonin, eds. Cold Spring Harb. Perspect. Biol. 6:a016154.

Goodenough, U. 2015. Historical perspective on Chlamydomonas as a model for basic research: 1950-1970. Plant J. 82: 365-369.

Umen, J., U. Goodenough, and J. Heitman. 2017. Eukaryotic sexual reproduction evoked “with a little help from my friends.” Cell 170: 1059-1061.

Goodenough, U., and B.D. Engle. 2023. Chapter 2: Cell Ultrastructure. In: In: Chlamydomonas Sourcebook, volume 1. U. Goodenough, ed. Amsterdam: Elsevier (in press).

Goodenough, U., and J-H. Lee. 2023. Chapter 3: Cell Walls. In: Chlamydomonas Sourcebook, volume 1. U. Goodenough, ed. Amsterdam: Elsevier (in press).

Goodenough, W.J. Snell, and J-H. Lee. 2023. Chapter 9: The Sexual Cycle. In: Chlamydomonas Sourcebook, volume 1. U. Goodenough, ed. Amsterdam: Elsevier (in press).

Public Outreach: The interface between science and religion (expanded at https://religious-naturalist-association.org/ursula-goodenough-2)

1989-       Council Member, Institute for Religion in an Age of Science (IRAS)

1992-96   President, IRAS

1992-       Member, Center for Advanced Study in Religion and Science 


1996-98   Member, Advisory Committee, AAAS Program of Dialogue Between

                Science and Religion

1996        IRAS Conference Co-Chair:  Epic of Evolution

1998        IRAS Conference Co-Chair: Interweaving Art, Science, and Spirituality

1998-       Editorial Advisory Board, Zygon: Journal of Religion and Science

2002-06   Vice-President for Development, IRAS

2003        IRAS Conference Co-Chair: Ecomorality

2006-08   Vice-President for Interdisciplinary Affairs, IRAS  

2007        IRAS Conference Co-Chair: Emergence: Nature’s Mode of Creativity

2009        IRAS Conference Co-Chair: Emergence: Nature’s Mode of Creativity –

                           The Human Dimension 

2014-       Religious Naturalist Association https://religious-naturalist-association.org, Co-Founder and



Publications in Public Outreach:

Goodenough, U.W. 1993. Creativity in science. Zygon 28:399-414.

Goodenough, U.W. 1994. What science can and cannot offer to a religious narrative. Zygon 29: 321-330.

Goodenough. U.W. 1994. The religious dimensions of the biological narrative. Zygon 29: 603-618.  

Goodenough, U.W. 1996. Biology: What one needs to know. Zygon 31:671-680.

Goodenough, U.W. 1998. The Sacred Depths of Nature. Oxford University Press (218 pp)

Goodenough, U.W. 2000. Reflections on science and technology.  Zygon 35: 1-8.

Goodenough, U.W. 2000.  Reflections on scientific and religious metaphor.  Zygon 35: 233-240.

Goodenough, U.W. 2000.  Religiopoiesis. Zygon 35: 352-355.

Goodenough, U.W. 2000.  Causality and subjectivity in the religious quest. Zygon 35: 725-734.

Goodenough, U.W. 2001.  Vertical and horizontal transcendence. Zygon 36: 21-31

Goodenough, U.W. 2001.  A setback to the dialogue: Response to Huston Smith. Zygon 36: 201-206.

Goodenough, U.W. 2001.  Genomes, Gould, and emergence. Zygon 36: 383-393.

Goodenough, U.W. and P. Woodruff.  2001.  Mindful virtue, mindful reverence. Zygon 36:585-595.

Goodenough, U.W. 2002.  How can scientific understandings of nature contribute to moral, spiritual, and religious wholeness and well being?  In: The Good in Nature and Humanity:  Connecting Science, Religion, and Spirit with the Natural World.  S.R. Kellert, T.J. Farnham, eds. Island Press 

Goodenough, U.W. 2003.  Religious naturalism and naturalizing morality. Zygon 38: 101-109.

Goodenough, U.W., and T.W. Deacon. 2003. From biology to consciousness to morality. Zygon 38: 801-819.

Goodenough, U.W. 2005.  Reductionism and holism, chance and selection, mechanism and mind. Zygon 40: 369-380.

Goodenough, U. and T.W. Deacon. 2007. The sacred emergence of nature.  In: The Oxford Handbook of Religion and Science.  P. Clayton, ed. Oxford University Press, pp 854-871.

Goodenough, U.W. 2007.  The emergence of sex.  Zygon 42: 857-872.

Rue, L. and U. Goodenough. 2009. A consilient curriculum. In: The Evolutionary Epic: Science’s Story and Humanity’s Response. C. Genet, R. Genet, B. Swimme, L. Palmer, L. Gibler, eds. Collins Foundation Press.

Goodenough, U. 2009. Ecomorality: Toward an ethic of sustainability. In: A Pivotal Moment: Population, justice, and the environmental challenge. L. Mazur, ed. Washington D.C., Island Press. pp 372-382.

Goodenough, U. 2011. The biological antecedents of human suffering. In: Routledge Companion to Religion and Science. J.W. Haag, G.P. Peterson, M.L. Spezio, eds. Routledge.

Goodenough, U. 2017. Who is a religious naturalist? Theol. and Sci. 15: 231-234.

Goodenough, U., M. Cavanaugh, and T. Macalister. 2018. Bringing religious naturalists together online. In: The Routledge Handbook of Religious Naturalism. D.A. Crosby and J.A. Stone, eds. Routledge. pp 310-316.

Goodenough, U. 2018. The pope and the religious naturalist: Our common ecomorality. In: Integral Ecology: Protecting our Common Home. G. Magill and J. Potter, eds. Cambridge Scholars Publishing. pp 10-25.

Goodenough, U.W., and J.E. Sherman. 2021. The emergence of selves and purpose. Zygon 56: 960-970

Goodenough, U.W. 2022. The Sacred Depths of Nature: How Life Has Emerged and Evolved, second edition. Oxford University Press (in press).    

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