Editing Arabidopsis thaliana (section)

==Use as a model organism==
{{Main|History of research on Arabidopsis thaliana|l1=History of research on ''Arabidopsis thaliana''}}
Botanists and biologists began to research ''A. thaliana'' in the early 1900s, and the first systematic description of mutants was done around 1945.<ref>[http://www.arabidopsis.org/portals/education/aboutarabidopsis.jsp] {{Webarchive|url=https://web.archive.org/web/20161022211543/http://www.arabidopsis.org/portals/education/aboutarabidopsis.jsp|date=22 October 2016}} TAIR: About ''[[Arabidopsis]]''</ref> ''A. thaliana'' is now widely used for studying [[plant sciences]], including [[genetics]], [[evolution]], population genetics, and plant development.<ref>{{cite journal |vauthors=Rensink WA, Buell CR |title=''Arabidopsis'' to rice. Applying knowledge from a weed to enhance our understanding of a crop species |journal=Plant Physiology |volume=135 |issue=2 |pages=622–9 |date=June 2004 |pmid=15208410 |pmc=514098 |doi=10.1104/pp.104.040170}}</ref><ref>{{cite journal |vauthors=Coelho SM, Peters AF, Charrier B, Roze D, Destombe C, Valero M, Cock JM |title=Complex life cycles of multicellular eukaryotes: new approaches based on the use of model organisms |journal=Gene |volume=406 |issue=1–2 |pages=152–70 |date=December 2007 |pmid=17870254 |doi=10.1016/j.gene.2007.07.025 |s2cid=24427325 |url=https://hal.archives-ouvertes.fr/hal-01926745/file/Life%20cycle%20review%20030807.pdf |access-date=29 June 2021 |archive-date=9 July 2021 |archive-url=https://web.archive.org/web/20210709181559/https://hal.archives-ouvertes.fr/hal-01926745/file/Life%20cycle%20review%20030807.pdf |url-status=live}}</ref><ref>{{cite journal |vauthors=Platt A, Horton M, Huang YS, Li Y, Anastasio AE, Mulyati NW, Agren J, Bossdorf O, Byers D, Donohue K, Dunning M, Holub EB, Hudson A, Le Corre V, Loudet O, Roux F, Warthmann N, Weigel D, Rivero L, Scholl R, Nordborg M, Bergelson J, Borevitz JO |title=The scale of population structure in ''Arabidopsis thaliana'' |journal=PLOS Genetics |volume=6 |issue=2 |pages=e1000843 |date=February 2010 |pmid=20169178 |pmc=2820523 |doi=10.1371/journal.pgen.1000843 |editor1-last=Novembre |editor1-first=John |doi-access=free }}</ref> Although ''A. thaliana'' the plant has little direct significance for agriculture, ''A. thaliana'' the model organism has revolutionized our understanding of the genetic, cellular, and molecular biology of flowering plants.
[[File:Arabidopsis mutants.jpg|thumb|right|A double-flower mutant, first documented in 1873]]

The first mutant in ''A. thaliana'' was documented in 1873 by [[Alexander Braun]], describing a [[double flower]] phenotype (the mutated gene was likely ''[[Agamous]]'', cloned and characterized in 1990).<ref>{{cite journal |vauthors=Yanofsky MF, Ma H, Bowman JL, Drews GN, Feldmann KA, Meyerowitz EM |title=The protein encoded by the ''Arabidopsis'' homeotic gene agamous resembles transcription factors |journal=Nature |volume=346 |issue=6279 |pages=35–9 |date=July 1990 |pmid=1973265 |doi=10.1038/346035a0 |bibcode=1990Natur.346...35Y |s2cid=4323431}}</ref> [[Friedrich Laibach]] (who had published the chromosome number in 1907) did not propose ''A. thaliana'' as a model organism, though, until 1943.<ref name="meyerowitz2001">{{cite journal |vauthors=Meyerowitz EM |title=Prehistory and history of ''Arabidopsis'' research |journal=Plant Physiology |volume=125 |issue=1 |pages=15–9 |date=January 2001 |pmid=11154286 |pmc=1539315 |doi=10.1104/pp.125.1.15}}</ref> His student, Erna Reinholz, published her thesis on ''A. thaliana'' in 1945, describing the first collection of ''A. thaliana'' mutants that they generated using [[X-ray]] [[mutagenesis]]. Laibach continued his important contributions to ''A. thaliana'' research by collecting a large number of accessions (often questionably referred to as "[[ecotype]]s"). With the help of Albert Kranz, these were organised into a large collection of 750 natural accessions of ''A. thaliana'' from around the world.

In the 1950s and 1960s, John Langridge and [[George Rédei]] played an important role in establishing ''A. thaliana'' as a useful organism for biological laboratory experiments. Rédei wrote several scholarly reviews instrumental in introducing the model to the scientific community. The start of the ''A. thaliana'' research community dates to a newsletter called ''Arabidopsis'' Information Service,<ref name="Arabidopsis-Information-Service">{{cite web |title=About AIS |website=[[The Arabidopsis Information Resource]] |date=2018-11-08 |url=http://www.arabidopsis.org/ais/newaishint.jsp |access-date=2021-04-25 |archive-date=27 April 2021 |archive-url=https://web.archive.org/web/20210427093913/https://www.arabidopsis.org/ais/newaishint.jsp |url-status=live}}</ref> established in 1964. The first International ''Arabidopsis'' Conference was held in 1965, in [[Göttingen]], Germany.

In the 1980s, ''A. thaliana'' started to become widely used in plant research laboratories around the world. It was one of several candidates that included maize, [[petunia]], and tobacco.<ref name="meyerowitz2001" /> The latter two were attractive, since they were easily transformable with the then-current technologies, while maize was a well-established genetic model for plant biology. The breakthrough year for ''A. thaliana'' as a model plant was 1986, in which [[Transfer DNA|T-DNA]]-mediated [[Transformation (genetics)#Plants|transformation]] and the first [[clone (genetics)|clone]]d ''A. thaliana'' gene were described.<ref>{{cite journal |vauthors=Lloyd AM, Barnason AR, Rogers SG, Byrne MC, Fraley RT, Horsch RB |title=Transformation of ''Arabidopsis thaliana'' with ''Agrobacterium tumefaciens'' |journal=Science |volume=234 |issue=4775 |pages=464–6 |date=October 1986 |pmid=17792019 |doi=10.1126/science.234.4775.464 |bibcode=1986Sci...234..464L |s2cid=22125701}}</ref><ref>{{cite journal |vauthors=Chang C, Meyerowitz EM |title=Molecular cloning and DNA sequence of the ''Arabidopsis thaliana'' alcohol dehydrogenase gene |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=83 |issue=5 |pages=1408–12 |date=March 1986 |pmid=2937058 |pmc=323085 |doi=10.1073/pnas.83.5.1408 |bibcode=1986PNAS...83.1408C |doi-access=free}}</ref>

===Genomics===

[[File:Plastomap of Arabidopsis thaliana.svg|thumb|Chloroplast genome map of ''A. thaliana'':<ref name="NCBI_NC_000932"/><ref name="Sato1999"/> Introns are in grey. Some genes consist of 5′ and 3′ portions. Strand 1 and 2 genes are transcribed clockwise and counterclockwise, respectively. The innermost circle provides the boundaries of the large and small single-copy regions (LSC and SSC, violet) separated by a pair of inverted repeats (IRa and IRB, black).]]

====Nuclear genome====

Due to the small size of its [[genome]], and because it is [[diploid]], ''Arabidopsis thaliana'' is useful for genetic mapping and [[sequencing]] — with about 157 megabase pairs<ref>{{cite journal |vauthors=Bennett MD, Leitch IJ, Price HJ, Johnston JS |title=Comparisons with ''Caenorhabditis'' (approximately 100 Mb) and ''Drosophila'' (approximately 175 Mb) using flow cytometry show genome size in ''Arabidopsis'' to be approximately 157 Mb, thus approximately 25% larger than the ''Arabidopsis'' genome initiative estimate of approximately 125 Mb |journal=Annals of Botany |volume=91 |issue=5 |pages=547–57 |date=April 2003 |pmid=12646499 |pmc=4242247 |doi=10.1093/aob/mcg057}}</ref> and five [[chromosome]]s, ''A. thaliana'' has one of the smallest genomes among plants.<ref name="Genome Assembly"/> It was long thought to have the smallest genome of all flowering plants,<ref>(Leutwileret al., 1984). In our survey Arabidopsis ...</ref> but that title is now considered to belong to plants in the genus ''[[Genlisea]]'', order [[Lamiales]], with ''[[Genlisea tuberosa]]'', a carnivorous plant, showing a genome size of approximately 61&nbsp;Mbp.<ref name="Fleischmann 2014">{{cite journal |vauthors=Fleischmann A, Michael TP, Rivadavia F, Sousa A, Wang W, Temsch EM, Greilhuber J, Müller KF, Heubl G |title=Evolution of genome size and chromosome number in the carnivorous plant genus Genlisea (Lentibulariaceae), with a new estimate of the minimum genome size in angiosperms |journal=Annals of Botany |volume=114 |issue=8 |pages=1651–63 |date=December 2014 |pmid=25274549 |pmc=4649684 |doi=10.1093/aob/mcu189}}</ref> It was the first plant genome to be sequenced, completed in 2000 by the ''Arabidopsis'' Genome Initiative.<ref>{{cite journal |author=The Arabidopsis Genome Initiative |title=Analysis of the genome sequence of the flowering plant ''Arabidopsis thaliana'' |journal=Nature |volume=408 |issue=6814 |pages=796–815 |date=December 2000 |pmid=11130711 |doi=10.1038/35048692 |bibcode=2000Natur.408..796T |doi-access=free}}</ref> The most up-to-date version of the ''A. thaliana'' genome is maintained by the Arabidopsis Information Resource.<ref>{{Cite web |title=TAIR - Genome Annotation |url=http://www.arabidopsis.org/portals/genAnnotation/gene_structural_annotation/annotation_data.jsp |access-date=29 December 2008 |archive-date=14 October 2008 |archive-url=https://web.archive.org/web/20081014183552/http://www.arabidopsis.org/portals/genAnnotation/gene_structural_annotation/annotation_data.jsp |url-status=live}}</ref>

The genome encodes ~27,600 [[protein]]-coding [[gene]]s and about 6,500 non-coding genes.<ref>{{Cite web |title=Details - Arabidopsis_thaliana - Ensembl Genomes 63 |url=http://ensembl.gramene.org/Arabidopsis_thaliana/Info/Annotation/#assembly |access-date=2021-06-15 |website=ensembl.gramene.org |language=en-gb |archive-date=24 June 2021 |archive-url=https://web.archive.org/web/20210624195651/http://ensembl.gramene.org/Arabidopsis_thaliana/Info/Annotation/#assembly |url-status=live}}</ref> However, the Uniprot database lists 39,342 proteins in their ''Arabidopsis'' reference proteome.<ref>{{Cite web |title=''Arabidopsis thaliana'' (Mouse-ear cress) |url=https://www.uniprot.org/proteomes/UP000006548 |access-date=2021-06-15 |website=www.uniprot.org |language=en |archive-date=21 May 2021 |archive-url=https://web.archive.org/web/20210521205551/https://www.uniprot.org/proteomes/UP000006548 |url-status=live}}</ref> Among the 27,600 protein-coding genes 25,402 (91.8%) are now annotated with "meaningful" product names,<ref>{{Cite journal |last1=Cheng |first1=Chia-Yi |last2=Krishnakumar |first2=Vivek |last3=Chan |first3=Agnes P. |last4=Thibaud-Nissen |first4=Françoise |last5=Schobel |first5=Seth |last6=Town |first6=Christopher D. |date=2017 |title=Araport11: a complete reannotation of the ''Arabidopsis thaliana'' reference genome |journal=The Plant Journal |language=en |volume=89 |issue=4 |pages=789–804 |doi=10.1111/tpj.13415 |pmid=27862469 |s2cid=12155857 |issn=1365-313X |doi-access=free }}</ref> although a large fraction of these proteins is likely only poorly understood and only known in general terms (e.g. as "DNA-binding protein without known specificity"). Uniprot lists more than 3,000 proteins as "uncharacterized" as part of the reference proteome.

====Chloroplast genome====

The plastome of ''A. thaliana'' is a 154,478 base-pair-long DNA molecule,<ref name="NCBI_NC_000932">{{cite web |url=https://www.ncbi.nlm.nih.gov/nuccore/NC_000932 |title=''Arabidopsis thaliana'' chloroplast, complete genome — NCBI accession number NC_000932.1 |publisher=National Center for Biotechnology Information |access-date=November 4, 2018 |archive-date=4 November 2018 |archive-url=https://web.archive.org/web/20181104211116/https://www.ncbi.nlm.nih.gov/nuccore/NC_000932 |url-status=live}}</ref> a size typically encountered in most flowering plants (see the [[List of sequenced plastomes#Flowering plants|list of sequenced plastomes]]). It comprises 136 genes coding for small subunit ribosomal proteins (''rps'', in yellow: see figure), large subunit ribosomal proteins (''rpl'', orange), hypothetical chloroplast open reading frame proteins (''ycf'', lemon), proteins involved in photosynthetic reactions (green) or in other functions (red), ribosomal RNAs (''rrn'', blue), and transfer RNAs (''trn'', black).<ref name="Sato1999">{{Cite journal |vauthors=Sato S, Nakamura Y, Kaneko T, Asamizu E, Tabata S |year=1999 |title=Complete structure of the chloroplast genome of ''Arabidopsis thaliana'' |journal=DNA Research |language=en |volume=6 |issue=5 |pages=283–290 |doi=10.1093/dnares/6.5.283 |pmid=10574454 |issn=1340-2838 |doi-access=free}}</ref>

====Mitochondrial genome====

The mitochondrial genome of ''A. thaliana'' is 367,808 base pairs long and contains 57 genes.<ref name="NCBI_BK010421">{{cite web |url=https://www.ncbi.nlm.nih.gov/nuccore/bk010421 |title=''Arabidopsis thaliana'' ecotype Col-0 mitochondrion, complete genome — NCBI accession number BK010421 |date=10 October 2018 |publisher=National Center for Biotechnology Information |access-date=April 10, 2019 |archive-date=12 April 2019 |archive-url=https://web.archive.org/web/20190412200017/https://www.ncbi.nlm.nih.gov/nuccore/bk010421 |url-status=live}}</ref> There are many repeated regions in the ''Arabidopsis'' mitochondrial genome. The largest repeats [[Genetic recombination|recombine]] regularly and isomerize the genome.<ref>{{cite journal |vauthors=Klein M, Eckert-Ossenkopp U, Schmiedeberg I, Brandt P, Unseld M, Brennicke A, Schuster W |year=1994 |title=Physical mapping of the mitochondrial genome of ''Arabidopsis thaliana'' by cosmid and YAC clones |journal=Plant Journal |volume=6 |issue=3 |pages=447–455 |doi=10.1046/j.1365-313X.1994.06030447.x |pmid=7920724 |doi-access=free}}</ref> Like most plant mitochondrial genomes, the ''Arabidopsis'' mitochondrial genome exists as a complex arrangement of overlapping branched and linear molecules ''in vivo''.<ref>{{cite journal |vauthors=Gualberto JM, Mileshina D, Wallet C, Niazi AK, Weber-Lotfi F, Dietrich A |year=2014 |title=The plant mitochondrial genome: dynamics and maintenance |journal=Biochimie |volume=100 |pages=107–120 |doi=10.1016/j.biochi.2013.09.016 |pmid=24075874}}</ref>

===Genetics===

[[Transformation (genetics)|Genetic transformation]] of ''A. thaliana'' is routine, using ''[[Agrobacterium tumefaciens]]'' to transfer [[deoxyribonucleic acid|DNA]] into the plant genome. The current protocol, termed "floral dip", involves simply dipping flowers into a solution containing ''Agrobacterium'' carrying a plasmid of interest and a detergent.<ref>{{cite journal |vauthors=Clough SJ, Bent AF |title=Floral dip: a simplified method for Agrobacterium-mediated transformation of ''Arabidopsis thaliana'' |journal=The Plant Journal |volume=16 |issue=6 |pages=735–43 |date=December 1998 |pmid=10069079 |doi=10.1046/j.1365-313x.1998.00343.x |s2cid=410286}}</ref><ref>{{cite journal |vauthors=Zhang X, Henriques R, Lin SS, Niu QW, Chua NH |title=Agrobacterium-mediated transformation of ''Arabidopsis thaliana'' using the floral dip method |journal=Nature Protocols |volume=1 |issue=2 |pages=641–6 |year=2006 |pmid=17406292 |doi=10.1038/nprot.2006.97 |s2cid=6906570}}</ref> This method avoids the need for [[tissue culture]] or plant regeneration.

The ''A. thaliana'' gene knockout collections are a unique resource for plant biology made possible by the availability of high-throughput transformation and funding for genomics resources. The site of T-DNA insertions has been determined for over 300,000 independent transgenic lines, with the information and seeds accessible through online T-DNA databases.<ref>{{Cite web |url=http://signal.salk.edu/cgi-bin/tdnaexpress |title=T-DNA Express: Arabidopsis Gene Mapping Tool |website=signal.salk.edu |access-date=19 October 2009 |archive-date=25 November 2009 |archive-url=https://web.archive.org/web/20091125151524/http://signal.salk.edu/cgi-bin/tdnaexpress/ |url-status=live}}</ref> Through these collections, insertional mutants are available for most genes in ''A. thaliana''.

Characterized accessions and mutant lines of ''A. thaliana'' serve as experimental material in laboratory studies. The most commonly used background lines are L''er'' (Landsberg ''erecta''), and Col, or Columbia.<ref name="arabidopsis.info">{{Cite web |url=http://arabidopsis.info/ |title=Eurasian Arabidopsis Stock Centre (uNASC) |website=arabidopsis.info |access-date=19 October 2009 |archive-date=12 December 2001 |archive-url=https://web.archive.org/web/20011212001728/http://arabidopsis.info/ |url-status=live}}</ref> Other background lines less-often cited in the scientific literature are Ws, or Wassilewskija, C24, Cvi, or Cape Verde Islands, Nossen, etc. (see for ex.<ref>{{cite journal |vauthors=Magliano TM, Botto JF, Godoy AV, Symonds VV, Lloyd AM, Casal JJ |title=New Arabidopsis recombinant inbred lines (Landsberg erecta x Nossen) reveal natural variation in phytochrome-mediated responses |journal=Plant Physiology |volume=138 |issue=2 |pages=1126–35 |date=June 2005 |pmid=15908601 |pmc=1150426 |doi=10.1104/pp.104.059071}}</ref>) Sets of closely related accessions named Col-0, Col-1, etc., have been obtained and characterized; in general, mutant lines are available through stock centers, of which best-known are the Nottingham Arabidopsis Stock Center-NASC<ref name="arabidopsis.info" /> and the Arabidopsis Biological Resource Center-ABRC in Ohio, USA.<ref>{{Cite web |url=https://abrc.osu.edu/ |title=ABRC |website=abrc.osu.edu |access-date=12 December 2020 |archive-date=25 February 2021 |archive-url=https://web.archive.org/web/20210225031213/https://abrc.osu.edu/ |url-status=live}}</ref>
The Col-0 accession was selected by Rédei from within a (nonirradiated) population of seeds designated 'Landsberg' which he received from Laibach.<ref>{{Cite web |url=http://arabidopsis.info/CollectionInfo?id=94 |title=NASC Collection Info |website=arabidopsis.info |access-date=15 February 2011 |archive-date=19 July 2011 |archive-url=https://web.archive.org/web/20110719022459/http://arabidopsis.info/CollectionInfo?id=94 |url-status=live}}</ref> Columbia (named for the location of Rédei's former institution, [[University of Missouri]]-[[Columbia, Missouri|Columbia]]) was the reference accession sequenced in the ''Arabidopsis'' Genome Initiative. The Later (Landsberg erecta) line was selected by Rédei (because of its short stature) from a Landsberg population he had mutagenized with X-rays. As the L''er'' collection of mutants is derived from this initial line, L''er''-0 does not correspond to the Landsberg accessions, which designated La-0, La-1, etc.

Trichome formation is initiated by the GLABROUS1 protein. [[Gene knockout|Knockouts]] of the corresponding gene lead to [[Glossary of botanical terms#glabrous|glabrous]] plants. This [[phenotype]] has already been used in [[genome editing|gene editing]] experiments and might be of interest as visual marker for plant research to improve gene editing methods such as [[CRISPR#Genome engineering|CRISPR/Cas9.]]<ref>{{cite journal |vauthors=Hahn F, Mantegazza O, Greiner A, Hegemann P, Eisenhut M, Weber AP |title=''Arabidopsis thaliana'' |language=en |journal=Frontiers in Plant Science |volume=8 |pages=39 |date=2017 |pmid=28174584 |pmc=5258748 |doi=10.3389/fpls.2017.00039 |doi-access=free}}</ref><ref>{{cite journal |vauthors=Hahn F, Eisenhut M, Mantegazza O, Weber AP |title=Arabidopsis With Cas9-Based Gene Targeting |journal=Frontiers in Plant Science |volume=9 |pages=424 |date=5 April 2018 |pmid=29675030 |pmc=5895730 |doi=10.3389/fpls.2018.00424 |doi-access=free}}</ref>

====Non-Mendelian inheritance controversy====
In 2005, scientists at [[Purdue University]] proposed that ''A. thaliana'' possessed an alternative to previously known mechanisms of [[DNA repair]], producing an unusual pattern of [[Mendelian inheritance|inheritance]], but the phenomenon observed (reversion of mutant copies of the ''[[HOTHEAD (gene)|HOTHEAD]]'' gene to a wild-type state) was later suggested to be an artifact because the mutants show increased outcrossing due to organ fusion.<ref>{{cite journal |vauthors=Lolle SJ, Victor JL, Young JM, Pruitt RE |title=Genome-wide non-mendelian inheritance of extra-genomic information in Arabidopsis |journal=Nature |volume=434 |issue=7032 |pages=505–9 |date=March 2005 |pmid=15785770 |doi=10.1038/nature03380 |bibcode=2005Natur.434..505L |s2cid=1352368}}[https://www.washingtonpost.com/wp-dyn/articles/A58349-2005Mar22_2.html Washington Post summary.] {{Webarchive |url=https://web.archive.org/web/20161118043014/http://www.washingtonpost.com/wp-dyn/articles/A58349-2005Mar22_2.html |date=18 November 2016 }}</ref><ref>{{cite journal |vauthors=Peng P, Chan SW, Shah GA, Jacobsen SE |title=Plant genetics: increased outcrossing in hothead mutants |journal=Nature |volume=443 |issue=7110 |pages=E8; discussion E8–9 |date=September 2006 |pmid=17006468 |doi=10.1038/nature05251 |bibcode=2006Natur.443E...8P |s2cid=4420979 |doi-access=free}}</ref><ref>{{cite journal |vauthors=Pennisi E |author-link=Elizabeth Pennisi |title=Genetics. Pollen contamination may explain controversial inheritance |journal=Science |volume=313 |issue=5795 |pages=1864 |date=September 2006 |pmid=17008492 |doi=10.1126/science.313.5795.1864 |s2cid=82215542|doi-access=free }}</ref>

===Lifecycle===

The plant's small size and rapid lifecycle are also advantageous for research. Having specialized as a [[spring ephemeral]], it has been used to found several laboratory strains that take about 6 weeks from germination to mature seed. The small size of the plant is convenient for cultivation in a small space, and it produces many seeds. Further, the selfing nature of this plant assists genetic experiments. Also, as an individual plant can produce several thousand seeds, each of the above criteria leads to ''A. thaliana'' being valued as a genetic model organism.

===Cellular biology===
''Arabidopsis'' is often the model for study of [[SNARE (protein)#In plants|SNAREs in plants]]. This has shown SNAREs to be heavily involved in [[vesicle trafficking]]. Zheng et al. 1999 found an ''Arabidopsis'' SNARE called {{visible anchor|AtVTI1a}} is probably essential to [[Golgi apparatus|Golgi]]-[[vacuole]] trafficking. This is still a wide open field and plant SNAREs' role in trafficking remains understudied.<ref name="Raikhel-2017">{{cite journal |last=Raikhel |first=Natasha V. |author-link=Natasha Raikhel |title=Firmly Planted, Always Moving |journal=[[Annual Review of Plant Biology]] |publisher=[[Annual Reviews (publisher)|Annual Reviews]] |volume=68 |issue=1 |date=2017-04-28 |issn=1543-5008 |doi=10.1146/annurev-arplant-042916-040829 |pages=1–27 |pmid=27860488|doi-access=free }}</ref>

===DNA repair===

The [[DNA]] of plants is vulnerable to [[ultraviolet]] light, and [[DNA repair]] mechanisms have evolved to avoid or repair genome damage caused by UV. Kaiser et al.<ref>Kaiser G, Kleiner O, Beisswenger C, Batschauer A. Increased DNA repair in Arabidopsis plants overexpressing CPD photolyase. Planta. 2009 Aug;230(3):505-15. doi: 10.1007/s00425-009-0962-y. Epub 2009 Jun 12. PMID 19521716</ref> showed that in ''A. thaliana'' cyclobutane pyrimidine dimers (CPDs) induced by UV light can be repaired by expression of CPD [[photolyase]].

===Germination in lunar regolith===
On May 12, 2022, [[NASA]] announced that specimens of ''Arabidopsis thaliana'' had been successfully germinated and grown in samples of [[lunar regolith]]. While the plants successfully germinated and grew into seedlings, they were not as robust as specimens that had been grown in [[volcanic ash]] as a control group, although the experiments also found some variation in the plants grown in regolith based on the location the samples were taken from, as ''A. thaliana'' grown in regolith gathered during [[Apollo 12]] & [[Apollo 17]] were more robust than those grown in samples taken during [[Apollo 11]].<ref>{{Cite web |last=Keeter |first=Bill |date=2022-05-12 |title=Scientists Grow Plants in Lunar Soil |url=http://www.nasa.gov/feature/biological-physical/scientists-grow-plants-in-soil-from-the-moon |access-date=2022-05-14 |website=NASA |archive-date=14 May 2022 |archive-url=https://web.archive.org/web/20220514200820/https://www.nasa.gov/feature/biological-physical/scientists-grow-plants-in-soil-from-the-moon/ |url-status=live }}</ref>