Susan Golden

Golden began her graduate career with Louis A. Sherman, where she used genetics to research the proteins in photosynthetic complexes of the cyanobacteria Synechoccus elongatus. Golden was the first to demonstrate that a mutant allele of the [https://www.wikigenes.org/e/gene/e/807514.html psbA gene] is sufficient to confer herbicide resistance in cyanobacteria. Other research later confirmed that this gene coded for a protein integral to the Photosystem II complex involved in photosynthesis.{{Cite journal|last1=Goussias|first1=Charilaos|last2=Boussac|first2=Alain|last3=Rutherford|first3=A William|date=2002-10-29|title=Photosystem II and photosynthetic oxidation of water: an overview.|journal=Philosophical Transactions of the Royal Society B: Biological Sciences|volume=357|issue=1426|pages=1369–1420|doi=10.1098/rstb.2002.1134|issn=0962-8436|pmc=1693055|pmid=12437876}} These findings also demonstrated that genetic manipulation of cyanobacteria was relatively simple, opening up S. elongatus as a model organism for future genetic experiments. During her postdoctoral research at the University of Chicago, in the lab of Robert Haselkorn, Golden continued to work on developing genetic manipulation techniques for Synechoccus elongatus in order to elucidate mechanisms of gene regulation in photosynthesis genes. In 1989, Golden's team discovered that the specific psbA allele expressed by cyanobacteria depended on the lighting conditions in which the colony was grown.{{Cite journal|last1=Schaefer|first1=M. R.|last2=Golden|first2=S. S.|date=1989-05-05|title=Light availability influences the ratio of two forms of D1 in cyanobacterial thylakoids.|journal=Journal of Biological Chemistry|language=en|volume=264|issue=13|pages=7412–7417|doi=10.1016/S0021-9258(18)83249-5|issn=0021-9258|pmid=2496127|doi-access=free}} This finding led her to investigate more generally how light influences expression of photosynthetic genes in the organism{{Cite web|url=http://www.nasonline.org/member-directory/members/2534843.html?referrer=https://www.google.com/|title=Susan Golden|website=www.nasonline.org|access-date=2017-04-12}} and contributes to the overall understanding of bacterial responses to environmental input.{{Cite journal|last1=Tandeau de Marsac|first1=Nicole|last2=Houmard|first2=Jean|date=1993-01-01|title=Adaptation of cyanobacteria to environmental stimuli: new steps towards molecular mechanisms|journal=FEMS Microbiology Letters|language=en|volume=104|issue=1–2|pages=119–189|doi=10.1111/j.1574-6968.1993.tb05866.x|issn=0168-6445|doi-access=free}} This line of inquiry necessitated the development of a technique for visualizing changes in gene expression in living organisms. During her time as a professor at the Texas A&M, Golden attempted to solve this problem by attaching a luciferase gene to the promoters of the cyanobacterial genes of interest and viewing the colonies with a night vision scope. The approach was a success, allowing for quantification of cyanobacterial gene expression in vivo over an extended time period. This technique drew the interest of chronobiologist Carl H. Johnson, with whom Golden would go on to collaborate and eventually discover the KaiABC complex .

Golden studies the endogenous rhythms of cyanobacteria, a group of photosynthetic bacteria known to have circadian clocks. She transformed Synechococcus elongatus, one of the better studied cyanobacteria species, with a luciferase reporter gene and showed circadian rhythm in bioluminescence. This reporter system was used to discover three key proteins related to the cyanobacterial clock: KaiA, KaiB, and KaiC. In collaboration with Carl H. Johnson and Takao Kondo, she demonstrated and described the molecular mechanism regulating circadian rhythms in S. elongatus PCC 7942,{{cite journal|last1=Cohen|first1=Susan|title=Circadian Rhythms in Cyanobacteria|journal=Microbiology and Molecular Biology Reviews|date=1 December 2015|volume=79|issue=4|pages=373–385|doi=10.1128/MMBR.00036-15|pmid=26335718|pmc=4557074}} the only model organism for a prokaryotic circadian clock. Susan Golden identified genes in the S. elongatus genome that contribute to circadian rhythm through mutational screens using transposons to disrupt genes and their function. In one mutation screen study, nineteen mutations were identified and mapped to the three kai genes; Inactivation of any single kai gene reduced kaiBC-promoter activity and abolished the circadian rhythm of expression of KaiA and KaiB.{{Cite journal|last1=Ishiura|first1=Masahiro|last2=Kutsuna|first2=Shinsuke|last3=Aoki|first3=Setsuyuki|last4=Iwasaki|first4=Hideo|last5=Andersson|first5=Carol R.|last6=Tanabe|first6=Akio|last7=Golden|first7=Susan S.|last8=Johnson|first8=Carl H.|last9=Kondo|first9=Takao|date=1998-09-04|title=Expression of a Gene Cluster kaiABC as a Circadian Feedback Process in Cyanobacteria|journal=Science|language=en|volume=281|issue=5382|pages=1519–1523|doi=10.1126/science.281.5382.1519|issn=0036-8075|pmid=9727980}}

S. elongatus has a circadian clock with an oscillator based only on three proteins: KaiA, KaiB, and KaiC. Circadium rhythm is generated based on KaiC phosphorylation and dephosphorylation. Light transmits energy and information to the cyanobacteria, affecting transcriptional regulation of the circadium clock. This 24-hour rhythm can be recreated in vitro with the addition of ATP.{{cite journal|last1=Dong|first1=Guogang|title=How a Cyanobacterium Tells Time|journal=Current Opinion in Microbiology|date=10 November 2008|volume=11|issue=6|pages=541–546|doi=10.1016/j.mib.2008.10.003|pmid=18983934|pmc=2692899}} The ratio of ATP/ADP fluctuates during the course of the day, and is sensed by KaiC, which phosphorylates or de-phosphorylates based on this signal. This Kai protein system is the simplest post-translational oscillator known so far.

In photosynthesizing cyanobacteria, the circadian clock is driven by light and reset by darkness. When Golden mutated the gene cikA, the clock was functional but could not be reset, resulting in the bacterial equivalent of permanent jet lag. CikA protein contains a domain that is structurally similar to KaiA, which was also found to be important in resetting the clock. CikA and KaiA bind to quinones, which carry electrons in the electron transport chain of photosynthesis. Quinones are oxidized in the dark and reduced in the light, and the redox state affects KaiA activity. When quinones are oxidized, KaiA separates from KaiC and binds to them, resetting the clock. Therefore, quinones are essential in transmitting light information to KaiC.

After moving to UC San Diego in 2008, Susan Golden's research converged with that of her husband, James Golden, to investigate biofuels. She currently researches the potential of utilizing cyanobacteria for industrial production of biofuels. Cyanobacteria are attractive due to simplistic genomes and ability to be genetically modified for industrial efficiency. Their photosynthetic nature may ideally be used to produce sustainably produce biofuels, potentially replacing the need for petroleum and other fossil fuels. They have simple requirements for growth, only requiring sunlight, water, and inorganic trace elements for fast growth. Cyanobacteria are capable of fixing atmospheric carbon (carbon dioxide) into bio-oils and biofuels.{{cite journal|last1=Sarsekeyeva|first1=Fariza|title=Cyanofuels: biofuels from cyanobacteria. Reality and perspectives|journal=Photosynthesis Research|date=22 February 2015|volume=125|issue=1|pages=325–340|doi=10.1007/s11120-015-0103-3|pmid=25702086|bibcode=2015PhoRe.125..329S |s2cid=15816714}}

In 2016, Golden and colleagues manually curated a model of metabolism in S. elongatus, indicating the importance of a linear tricarboxylic acid cycle (TCA) metabolic pathway and potential modifications for bio-industrial application.{{Cite journal|last1=Broddrick|first1=Jared T.|last2=Rubin|first2=Benjamin E.|last3=Welkie|first3=David G.|last4=Du|first4=Niu|last5=Mih|first5=Nathan|last6=Diamond|first6=Spencer|last7=Lee|first7=Jenny J.|last8=Golden|first8=Susan S.|last9=Palsson|first9=Bernhard O.|date=2016-12-20|title=Unique attributes of cyanobacterial metabolism revealed by improved genome-scale metabolic modeling and essential gene analysis|journal=Proceedings of the National Academy of Sciences|language=en|volume=113|issue=51|pages=E8344–E8353|doi=10.1073/pnas.1613446113|issn=0027-8424|pmc=5187688|pmid=27911809|bibcode=2016PNAS..113E8344B |doi-access=free}}

• National Science Foundation Presidential Young Investigator Award, 1989 - 1995
• Fellow of the American Academy of Microbiology, elected in 2000
• Texas A&M Distinguished Professor, 2003
• Member of the Faculty of 1000 Biology, 2008 - 2015{{Cite web |title=Susan Golden {{!}} Faculty Member {{!}} Faculty Opinions |url=https://facultyopinions.com/prime/thefaculty/member/2775310791432883 |access-date=2022-03-16 |website=facultyopinions.com}}
• UC San Diego Distinguished Professor and Chancellor’s Associates Chair in Molecular Biology, 2008–present
• Member of the National Academy of Sciences, elected in 2010{{Cite web |title=Susan S. Golden |url=http://www.nasonline.org/member-directory/members/2534843.html |access-date=2022-03-16 |website=www.nasonline.org}}
• Howard Hughes Medical Institute Professor, 2014–present
• Aschoff and Honma Prize for Biological Rhythm Research, 2018{{Cite web |title=Susan Golden to Receive International Research Honor |url=https://biology.ucsd.edu/about/news/article_030218.html |access-date=2022-03-16 |website=biology.ucsd.edu}}

{{excessive examples|section|date=March 2022}}

• {{cite journal

|author=Kondo T, Strayer CA, Kulkarni RD, Taylor W, Ishiura M, Golden SS, Johnson CH

|title=Circadian rhythms in prokaryotes: luciferase as a reporter of circadian gene expression in cyanobacteria

|journal=Proc Natl Acad Sci USA

|volume=90

|issue=12

|pages=5672–6

|year=1993

|pmid=8516317

|pmc=46783

|doi=10.1073/pnas.90.12.5672

|bibcode=1993PNAS...90.5672K

|doi-access=free

}}

• {{cite journal

|author=Kondo T, Tsinoremas NF, Golden SS, Johnson CH, Kutsuna S, Ishiura M

|title=Circadian clock mutants of cyanobacteria

|journal=Science

|volume=266

|issue=5188

|pages=1233–6

|year=1994

|pmid=7973706

|doi=10.1126/science.7973706

|bibcode=1994Sci...266.1233K

}}

• {{cite journal

|author=Liu Y, Golden SS, Kondo T, Ishiura M, Johnson CH

|title=Bacterial luciferase as a reporter of circadian gene expression in cyanobacteria

|journal=J Bacteriol

|volume=177

|issue=8

|pages=2080–6

|year=1995

|pmid=7536731

|pmc=176852

|doi=10.1128/jb.177.8.2080-2086.1995

}}

• {{cite journal

|author=Golden SS

|title=Light-responsive gene expression in cyanobacteria

|journal=J Bacteriol

|volume=177

|issue=7

|pages=1651–4

|year=1995

|pmid=7896684

|pmc=176789

|doi=10.1128/jb.177.7.1651-1654.1995

}}

• {{cite journal

|author=Liu Y, Tsinoremas NF, Johnson CH, Lebedeva NV, Golden SS, Ishiura M, Kondo T

|title=Circadian orchestration of gene expression in cyanobacteria

|journal=Genes Dev

|volume=9

|issue=12

|pages=1469–78

|year=1995

|pmid=7601351

|doi=10.1101/gad.9.12.1469

|doi-access=free

}}

• {{cite journal

|author=Tsinoremas NF, Ishiura M, Kondo T, Andersson CR, Tanaka K, Takahashi H, Johnson CH, Golden SS

|title=A sigma factor that modifies the circadian expression of a subset of genes in cyanobacteria

|journal=EMBO J

|volume=15

|issue=10

|pages=2488–95

|year=1996

|pmid=8665856

|pmc=450181

|doi=10.1002/j.1460-2075.1996.tb00606.x

}}

• {{cite journal

|author=Johnson CH, Golden SS, Ishiura M, Kondo T

|title=Circadian clocks in prokaryotes

|journal=Mol Microbiol

|volume=21

|issue=1

|pages=5–11

|year=1996

|pmid=8843429

|doi=10.1046/j.1365-2958.1996.00613.x

|s2cid=40431382

}}

• {{cite journal

|author=Kondo T, Mori T, Lebedeva NV, Aoki S, Ishiura M, Golden SS

|title=Circadian rhythms in rapidly dividing cyanobacteria

|journal=Science

|volume=275

|issue=5297

|pages=224–7

|year=1997

|pmid=8985018

|doi=10.1126/science.275.5297.224

|s2cid=31261881

}}

• {{cite journal

|author=Golden SS, Ishiura M, Johnson CH, Kondo T

|title=Cyanobacterial circadian rhythms

|journal=Annu Rev Plant Physiol Plant Mol Biol

|volume=48

|pages=327–354

|year=1997

|pmid=15012266

|doi=10.1146/annurev.arplant.48.1.327

}}

• {{cite journal

|author=Ouyang Y, Andersson CR, Kondo T, Golden SS, Johnson CH

|title=Resonating circadian clocks enhance fitness in cyanobacteria

|journal=Proc Natl Acad Sci USA

|volume=95

|issue=15

|pages=8660–4

|year=1998

|pmid=9671734

|pmc=21132

|doi=10.1073/pnas.95.15.8660

|bibcode=1998PNAS...95.8660O

|doi-access=free

}}

• {{cite journal

|author=Ishiura M, Kutsuna S, Aoki S, Iwasaki H, Andersson CR, Tanabe A, Golden SS, Johnson CH, Kondo T

|title=Expression of a gene cluster kaiABC as a circadian feedback process in cyanobacteria

|journal=Science

|volume=281

|issue=5382

|pages=1519–23

|year=1998

|pmid=9727980

|doi=10.1126/science.281.5382.1519

}}

• {{cite journal

|author=Johnson CH, Golden SS, Kondo T

|title=Adaptive significance of circadian programs in cyanobacteria

|journal=Trends Microbiol

|volume=6

|issue=10

|pages=407–10

|year=1998

|pmid=9807785

|doi=10.1016/s0966-842x(98)01356-0

}}

• {{cite journal

|author=Golden SS, Johnson CH, Kondo T

|title=The cyanobacterial circadian system: a clock apart

|journal=Curr Opin Microbiol

|volume=1

|issue=6

|pages=669–73

|year=1998

|pmid=10066545

|doi=10.1016/s1369-5274(98)80113-6

}}

• {{cite journal

|author=Katayama M, Tsinoremas NF, Kondo T, Golden SS

|title=cpmA, a gene involved in an output pathway of the cyanobacterial circadian system

|journal=J Bacteriol

|volume=181

|issue=11

|pages=3516–24

|year=1999

|pmid=10348865

|pmc=93820

|doi=10.1128/JB.181.11.3516-3524.1999

}}

• {{cite journal

|author=Johnson CH, Golden SS

|title=Circadian programs in cyanobacteria: adaptiveness and mechanism

|journal=Annu Rev Microbiol

|volume=53

|pages=389–409

|year=1999

|pmid=10547696

|doi=10.1146/annurev.micro.53.1.389

}}

• {{cite book

|author=Andersson CR, Tsinoremas NF, Shelton J, Lebedeva NV, Yarrow J, Min H, Golden SS

|title=Bioluminescence and Chemiluminescence Part C

|chapter=Application of bioluminescence to the study of circadian rhythms in cyanobacteria

|volume=305

|pages=527–42

|year=2000

|pmid=10812624

|doi=10.1016/s0076-6879(00)05511-7

|series=Methods in Enzymology

|isbn=9780121822064

}}

• {{cite journal

|author=Williams SB, Vakonakis I, Golden SS, LiWang AC

|title=Structure and function from the circadian clock protein KaiA of Synechococcus elongatus: a potential clock input mechanism

|journal=Proc Natl Acad Sci USA

|volume=99

|issue=24

|pages=15357–62

|year=2002

|pmid=12438647

|pmc=137721

|doi=10.1073/pnas.232517099

|bibcode=2002PNAS...9915357W

|doi-access=free

}}

• {{cite journal

|author=Ditty JL, Williams SB, Golden SS

|title=A cyanobacterial circadian timing mechanism

|journal=Annu Rev Genet

|volume=37

|pages=513–43

|year=2003

|pmid=14616072

|doi=10.1146/annurev.genet.37.110801.142716

|s2cid=36703896

}}

• {{cite journal

|author=Golden SS

|title=Timekeeping in bacteria: the cyanobacterial circadian clock

|journal=Curr Opin Microbiol

|volume=6

|issue=6

|pages=535–40

|year=2003

|pmid=14662347

|doi=10.1016/j.mib.2003.10.012

}}

• {{cite journal

|author=Golden SS

|title=Meshing the gears of the cyanobacterial circadian clock

|journal=Proc Natl Acad Sci USA

|volume=101

|issue=38

|pages=13697–8

|year=2004

|pmid=15367731

|pmc=518819

|doi=10.1073/pnas.0405623101

|bibcode=2004PNAS..10113697G

|doi-access=free

}}

• {{cite journal

|author=Mackey SR, Golden SS

|title=Winding up the cyanobacterial circadian clock

|journal=Trends Microbiol

|volume=15

|issue=9

|pages=381–8

|year=2007

|pmid=17804240

|doi=10.1016/j.tim.2007.08.005

}}

• {{cite journal

|author=Kim YI, Dong G, Carruthers CW Jr, Golden SS, LiWang A

|title=The day/night switch in KaiC, a central oscillator component of the circadian clock of cyanobacteria

|journal=Proc Natl Acad Sci USA

|volume=105

|issue=35

|pages=12825–30

|year=2008

|pmid=18728181

|doi=10.1073/pnas.0800526105

|pmc=2529086

|bibcode=2008PNAS..10512825K

|doi-access=free

}}

• {{cite journal

|author=Dong G, Golden SS

|title=How a cyanobacterium tells time

|journal=Curr Opin Microbiol

|volume=11

|issue=6

|pages=541–6

|year=2008

|pmid=18983934

|doi=10.1016/j.mib.2008.10.003

|pmc=2692899

}}

• {{cite journal

|author=Chen Y, Kim YI, Mackey SR, Holtman CK, Liwang A, Golden SS

|title=A novel allele of kaiA shortens the circadian period and strengthens interaction of oscillator components in the cyanobacterium Synechococcus elongatus PCC 7942

|journal=J Bacteriol

|volume=191

|issue=13

|pages=4392–400

|year=2009

|pmid=19395479

|pmc=2698500

|doi=10.1128/JB.00334-09

}}

• {{cite journal

|author=Dong G, Yang Q, Wang Q, Kim YI, Wood TL, Osteryoung KW, van Oudenaarden A, Golden SS

|title=Elevated ATPase activity of KaiC applies a circadian checkpoint on cell division in Synechococcus elongatus

|journal=Cell

|volume=140

|issue=4

|pages=529–39

|year=2010

|pmid=20178745

|pmc=3031423

|doi=10.1016/j.cell.2009.12.042

}}

• {{cite journal

|author=Dong G, Kim YI, Golden SS

|title=Simplicity and complexity in the cyanobacterial circadian clock mechanism

|journal=Curr Opin Genet Dev

|volume=20

|issue=6

|pages=619–25

|year=2010

|pmid=20934870

|pmc=2982900

|doi=10.1016/j.gde.2010.09.002

}}

• {{cite journal

|author=Cohen SE, Golden SS

|title=Circadian Rhythms in Cyanobacteria

|journal=Microbiol Mol Biol Rev

|volume=79

|issue=4

|pages=373–85

|year=2015

|pmid=26335718

|pmc=4557074

|doi=10.1128/MMBR.00036-15

}}

• {{cite journal

|author=Rubin BE, Wetmore KM, Price MN, Diamond S, Shultzaberger RK, Lowe LC, Curtin G, Arkin AP, Deutschbauer A, Golden SS

|title=The essential gene set of a photosynthetic organism

|journal=Proc Natl Acad Sci USA

|volume=112

|issue=48

|pages=E6634–43

|year=2015

|pmid=26508635

|pmc=4672817

|doi=10.1073/pnas.1519220112

|bibcode=2015PNAS..112E6634R

|doi-access=free

}}

• {{cite journal

|author=Tseng R, Goularte NF, Chavan A, Luu J, Cohen SE, Chang YG, Heisler J, Li S, Michael AK, Tripathi S, Golden SS, LiWang A, Partch CL

|title=Structural basis of the day-night transition in a bacterial circadian clock

|journal=Science

|volume=355

|issue=6330

|pages=1174–1180

|year=2017

|pmid=28302851

|pmc=5441561

|doi=10.1126/science.aag2516

|bibcode=2017Sci...355.1174T

}}

{{reflist}}

{{Authority control}}

{{DEFAULTSORT:Golden, Susan}}

Category:Living people

Category:American chronobiologists

Category:Members of the United States National Academy of Sciences

Category:Year of birth missing (living people)

Category:Fellows of the American Academy of Microbiology

Category:American women molecular biologists

Category:21st-century American women academics

Category:Mississippi University for Women alumni

Category:People from Pine Bluff, Arkansas

Category:Scientists from Arkansas

Category:University of California, San Diego faculty

Category:University of Missouri alumni

Category:Texas A&M University faculty

Category:University of Chicago faculty