A. Thaliana
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''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A
winter annual An annual plant is a plant that completes its biological life cycle, life cycle, from germination to the production of seeds, within one growing season, and then dies. The length of growing seasons and period in which they take place vary accor ...
with a relatively short lifecycle, ''A. thaliana'' is a popular
model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...
in plant biology and genetics. For a complex multicellular
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
, ''A. thaliana'' has a relatively small genome around 135 mega
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
s. It was the first plant to have its genome sequenced, and is a popular tool for understanding the molecular biology of many plant traits, including flower development and light sensing.


Description

''Arabidopsis thaliana'' is an annual (rarely biennial) plant, usually growing to 20–25 cm tall. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
form a rosette at the base of the plant, with a few leaves also on the flowering stem. The basal leaves are green to slightly purplish in color, 1.5–5 cm long, and 2–10 mm broad, with an entire to coarsely serrated margin; the stem leaves are smaller and unstalked, usually with an entire margin. Leaves are covered with small, unicellular hairs called trichomes. The flowers are 3 mm in diameter, arranged in a corymb; their structure is that of the typical Brassicaceae. The fruit is a siliqua 5–20 mm long, containing 20–30 seeds.Flora of NW Europe
''Arabidopsis thaliana''
Blamey, M. & Grey-Wilson, C. (1989). ''Flora of Britain and Northern Europe''. Flora of Pakistan
''Arabidopsis thaliana''
Flora of China
''Arabidopsis thaliana''
Roots are simple in structure, with a single primary root that grows vertically downward, later producing smaller lateral roots. These roots form interactions with rhizosphere bacteria such as '' Bacillus megaterium''. ''A. thaliana'' can complete its entire lifecycle in six weeks. The central stem that produces flowers grows after about 3 weeks, and the flowers naturally self-pollinate. In the lab, ''A. thaliana'' may be grown in Petri plates, pots, or hydroponics, under fluorescent lights or in a greenhouse.


Taxonomy

The plant was first described in 1577 in the Harz Mountains by (1542–1583), a physician from
Nordhausen Nordhausen may refer to: * Nordhausen (district), a district in Thuringia, Germany ** Nordhausen, Thuringia, a city in the district **Nordhausen station, the railway station in the city * Nordhouse, a commune in Alsace (German: Nordhausen) * Narost ...
, Thüringen, Germany, who called it ''Pilosella siliquosa''. In 1753, Carl Linnaeus renamed the plant ''Arabis thaliana'' in honor of Thal. In 1842, German botanist Gustav Heynhold erected the new genus ''Arabidopsis'' and placed the plant in that genus. The generic name, ''
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organi ...
'', comes from Greek, meaning "resembling '' Arabis''" (the genus in which Linnaeus had initially placed it). Thousands of natural inbred accessions of ''A. thaliana'' have been collected from throughout its natural and introduced range. These accessions exhibit considerable genetic and phenotypic variation, which can be used to study the adaptation of this species to different environments.


Distribution and habitat

''A. thaliana'' is native to Europe, Asia, and Africa, and its geographic distribution is rather continuous from the Mediterranean to Scandinavia and Spain to Greece. It also appears to be native in tropical alpine ecosystems in Africa and perhaps South Africa. It has been introduced and naturalized worldwide, including in North America around the 17th century. ''A. thaliana'' readily grows and often pioneers rocky, sandy, and calcareous soils. It is generally considered a weed, due to its widespread distribution in agricultural fields, roadsides, railway lines, waste ground, and other disturbed habitats, but due to its limited competitive ability and small size, it is not categorized as a noxious weed. Like most Brassicaceae species, ''A. thaliana'' is edible by humans in a salad or cooked, but it does not enjoy widespread use as a spring vegetable.


Use as a model organism

Botanists and biologists began to research ''A. thaliana'' in the early 1900s, and the first systematic description of mutants was done around 1945. ''A. thaliana'' is now widely used for studying plant sciences, including genetics, evolution, population genetics, and plant development. Although ''A. thaliana'' has little direct significance for agriculture, several of its traits make it a useful model for understanding the genetic, cellular, and molecular biology of flowering plants. The first mutant in ''A. thaliana'' was documented in 1873 by Alexander Braun, describing a
double flower "Double-flowered" describes varieties of flowers with extra petals, often containing flowers within flowers. The double-flowered trait is often noted alongside the scientific name with the abbreviation ''fl. pl.'' (''flore pleno'', a Latin ablati ...
phenotype (the mutated gene was likely '' Agamous'', cloned and characterized in 1990).
Friedrich Laibach Friedrich Laibach (born April 2, 1885, in Limburg, Germany) was a German botanist. Life and work Laibach was promoted in 1907 at the University of Bonn (with Eduard Strasburger). In 1919 he qualified as a professor at the University of Frankfurt (b ...
(who had published the chromosome number in 1907) did not propose ''A. thaliana'' as a model organism, though, until 1943. 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 " ecotypes"). 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 George P. Rédei (June 14, 1921 – November 10, 2008) was a Hungarian plant biologist, professor, author and member of the Hungarian Academy of Sciences. Biography George Rédei was born June 14, 1921, in Vienna, Austria, to Kalman and Margit ...
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, 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. 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 T-DNA-mediated transformation and the first
clone Clone or Clones or Cloning or Cloned or The Clone may refer to: Places * Clones, County Fermanagh * Clones, County Monaghan, a town in Ireland Biology * Clone (B-cell), a lymphocyte clone, the massive presence of which may indicate a pathologi ...
d ''A. thaliana'' gene were described.


Genomics


Nuclear genome

Due to the small size of its genome, and because it is
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, ''Arabidopsis thaliana'' is useful for genetic mapping and
sequencing In genetics and biochemistry, sequencing means to determine the primary structure (sometimes incorrectly called the primary sequence) of an unbranched biopolymer. Sequencing results in a symbolic linear depiction known as a sequence which succ ...
— with about 157 megabase pairs and five chromosomes, ''A. thaliana'' has one of the smallest genomes among plants. It was long thought to have the smallest genome of all flowering plants, but that title is now considered to belong to plants in the genus ''
Genlisea ''Genlisea'' ( ) is a genus of carnivorous plants also known as corkscrew plants. The 30 or so species grow in wet terrestrial to semi-aquatic environments distributed throughout Africa and Central and South America. The plants use highly modifi ...
'', order Lamiales, with ''
Genlisea tuberosa ''Genlisea tuberosa'' is a carnivorous species in the genus '' Genlisea'' (family Lentibulariaceae) that is endemic to Brazil and found only in '' campos rupestres'' vegetation. Lacking any roots, it has unpigmented bundles of "rootlike" subterr ...
'', a carnivorous plant, showing a genome size of approximately 61 Mbp. It was the first plant genome to be sequenced, completed in 2000 by the Arabidopsis Genome Initiative. The most up-to-date version of the ''A. thaliana'' genome is maintained by the Arabidopsis Information Resource. The genome encodes ~27,600 protein-coding genes and about 6,500 non-coding genes. However, the Uniprot database lists 39,342 proteins in their ''Arabidopsis'' reference proteome. Among the 27,600 protein-coding genes 25,402 (91.8%) are now annotated with "meaningful" product names, 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, a size typically encountered in most flowering plants (see the 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).


Mitochondrial genome

The mitochondrial genome of ''A. thaliana'' is 367,808 base pairs long and contains 57 genes. There are many repeated regions in the ''Arabidopsis'' mitochondrial genome. The largest repeats recombine regularly and isomerize the genome. Like most plant mitochondrial genomes, the ''Arabidopsis'' mitochondrial genome exists as a complex arrangement of overlapping branched and linear molecules ''in vivo''.


Genetics

Genetic transformation In molecular biology and genetics, transformation is the genetic alteration of a cell resulting from the direct uptake and incorporation of exogenous genetic material from its surroundings through the cell membrane(s). For transformation to ta ...
of ''A. thaliana'' is routine, using ''
Agrobacterium tumefaciens ''Agrobacterium radiobacter'' (more commonly known as ''Agrobacterium tumefaciens'') is the causal agent of crown gall disease (the formation of tumours) in over 140 species of eudicots. It is a rod-shaped, Gram-negative soil bacterium. Sympto ...
'' to transfer 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. 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. 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. 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.) 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 and the Arabidopsis Biological Resource Center-ABRC in Ohio, USA. The Col-0 accession was selected by Rédei from within a (nonirradiated) population of seeds designated 'Landsberg' which he received from Laibach. Columbia (named for the location of Rédei's former institution, University of Missouri-
Columbia Columbia may refer to: * Columbia (personification), the historical female national personification of the United States, and a poetic name for America Places North America Natural features * Columbia Plateau, a geologic and geographic region in ...
) 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.
Knockouts Knockouts and Knockouts Haircuts for Men are the trade names of a privately held American salon chain (international as of early 2011), Knockouts LLC, based in Irving, Texas. Knockouts are full-service grooming salons with a boxing (and mixed m ...
of the corresponding gene lead to glabrous plants. This phenotype has already been used in gene editing experiments and might be of interest as visual marker for plant research to improve gene editing methods such as CRISPR/Cas9.


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 inheritance, but the phenomenon observed (reversion of mutant copies of the ''
HOTHEAD Hothead may refer to: * "Hothead" (''Courage the Cowardly Dog''), a 1999 television episode * ''Hothead'' (film), a 1979 French film * HOTHEAD (gene), a gene in ''Arabidopsis thaliana'' that encodes a flavin adenine dinucleotide-containing oxido ...
'' gene to a wild-type state) was later suggested to be an artifact because the mutants show increased outcrossing due to organ fusion.


Lifecycle

The plant's small size and rapid lifecycle are also advantageous for research. Having specialized as a
spring ephemeral An ephemeral plant is one marked by short life cycles. The word ephemeral means transitory or quickly fading. In regard to plants, it refers to several distinct growth strategies. The first, spring ephemeral, refers to perennial plants that emerge ...
, 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
SNAREs in plants SNARE proteins – " SNAP REceptor" – are a large protein family consisting of at least 24 members in yeasts, more than 60 members in mammalian cells, and some numbers in plants. The primary role of SNARE proteins is to mediate vesicle fu ...
. This has shown SNAREs to be heavily involved in
vesicle trafficking In cell biology, a vesicle is a structure within or outside a cell, consisting of liquid or cytoplasm enclosed by a lipid bilayer. Vesicles form naturally during the processes of secretion (exocytosis), uptake (endocytosis) and transport of mater ...
. Zheng et al. 1999 found an ''Arabidopsis'' SNARE called is probably essential to Golgi- vacuole trafficking. This is still a wide open field and plant SNAREs' role in trafficking remains understudied.


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. 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 12 (November 14–24, 1969) was the sixth crewed flight in the United States Apollo program and the second to land on the Moon. It was launched on November 14, 1969, by NASA from the Kennedy Space Center, Florida. Commander Pete Conra ...
&
Apollo 17 Apollo 17 (December 7–19, 1972) was the final mission of NASA's Apollo program, the most recent time humans have set foot on the Moon or traveled beyond low Earth orbit. Commander Gene Cernan and Lunar Module Pilot Harrison Schmitt walked on ...
were more robust than those grown in samples taken during Apollo 11.


Development


Flower development

''A. thaliana ''has been extensively studied as a model for flower development. The developing flower has four basic organs - sepals,
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s,
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s, and carpels (which go on to form
pistil Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
s). These organs are arranged in a series of whorls, four sepals on the outer whorl, followed by four petals inside this, six stamens, and a central carpel region. Homeotic mutations in ''A. thaliana'' result in the change of one organ to another—in the case of the ''agamous'' mutation, for example, stamens become petals and carpels are replaced with a new flower, resulting in a recursively repeated sepal-petal-petal pattern. Observations of homeotic mutations led to the formulation of the
ABC model of flower development The ABC model of flower development is a scientific model of the process by which flowering plants produce a pattern of gene expression in meristems that leads to the appearance of an organ oriented towards sexual reproduction, a flower. There a ...
by E. Coen and E. Meyerowitz. According to this model, floral organ identity genes are divided into three classes - class A genes (which affect sepals and petals), class B genes (which affect petals and stamens), and class C genes (which affect stamens and carpels). These genes code for transcription factors that combine to cause tissue specification in their respective regions during development. Although developed through study of ''A. thaliana'' flowers, this model is generally applicable to other flowering plants.


Leaf development

Studies of ''A. thaliana'' have provided considerable insights with regards to the genetics of leaf morphogenesis, particularly in dicotyledon-type plants. Much of the understanding has come from analyzing mutants in leaf development, some of which were identified in the 1960s, but were not analysed with genetic and molecular techniques until the mid-1990s. ''A. thaliana'' leaves are well suited to studies of leaf development because they are relatively simple and stable. Using ''A. thaliana'', the genetics behind leaf shape development have become more clear and have been broken down into three stages: The initiation of the
leaf primordium A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
, the establishment of dorsiventrality, and the development of a marginal meristem. Leaf primordia are initiated by the suppression of the genes and proteins of class I '' KNOX'' family (such as ''SHOOT APICAL MERISTEMLESS''). These class I KNOX proteins directly suppress gibberellin biosynthesis in the leaf primordium. Many genetic factors were found to be involved in the suppression of these class I ''KNOX'' genes in leaf primordia (such as ''ASYMMETRIC LEAVES1,'' ''BLADE-ON-PETIOLE1'', ''SAWTOOTH1'', etc.). Thus, with this suppression, the levels of gibberellin increase and leaf primordium initiate growth. The establishment of leaf dorsiventrality is important since the dorsal (adaxial) surface of the leaf is different from the ventral (abaxial) surface.


Microscopy

''A. thaliana'' is well suited for
light microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...
analysis. Young seedlings on the whole, and their roots in particular, are relatively translucent. This, together with their small size, facilitates live cell imaging using both fluorescence and confocal laser scanning microscopy. By wet-mounting seedlings in water or in culture media, plants may be imaged uninvasively, obviating the need for fixation and sectioning and allowing time-lapse measurements. Fluorescent protein constructs can be introduced through transformation. The developmental stage of each cell can be inferred from its location in the plant or by using fluorescent protein
markers The term Marker may refer to: Common uses * Marker (linguistics), a morpheme that indicates some grammatical function * Marker (telecommunications), a special-purpose computer * Boundary marker, an object that identifies a land boundary * Marke ...
, allowing detailed developmental analysis.


Physiology


Light sensing, light emission, and circadian biology

The photoreceptors phytochromes A, B, C, D, and E mediate red light-based phototropic response. Understanding the function of these receptors has helped plant biologists understand the signaling cascades that regulate photoperiodism,
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
,
de-etiolation Etiolation is a process in flowering plants grown in partial or complete absence of light. It is characterized by long, weak stems; smaller leaves due to longer internodes; and a pale yellow color (chlorosis). The development of seedlings in t ...
, and
shade avoidance Shade avoidance is a set of responses that plants display when they are subjected to the shade of another plant. It often includes elongation, altered flowering time, increased apical dominance and altered partitioning of resources. This set of r ...
in plants. The genes ''
FCA FCA may refer to: Arts * Federation of Canadian Artists * Foundation for Contemporary Art, in Ghana * Foundation for Contemporary Arts, in the United States Business and economics * False Claims Act, a United States federal law * Federal Cus ...
'', '' fy'', ''
fpa FPA may refer to: Broadcasting and entertainment * ''Fancy Pants Adventures'', an online game * Feminist Porn Award, a Canadian adult film award * First-person adventure, a video game genre * Fundação Padre Anchieta, a Brazilian educational me ...
'', '' LUMINIDEPENDENS'' (''ld''), ''
fly Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
'', '' fve'' and ''
FLOWERING LOCUS C Flowering Locus C (''FLC'') is a MADS-box gene that in late-flowering ecotypes of the plant '' Arabidopsis thaliana'' is responsible for vernalization. In a new seedling ''FLC'' is expressed, which prevents flowering. Upon exposure to cold, less ...
'' (''FLC'') are involved in
photoperiod Photoperiodism is the physiological reaction of organisms to the length of night or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light a ...
triggering of flowering and vernalization. Specifically Lee et al 1994 find ''ld'' produces a homeodomain and Blazquez et al 2001 that ''fve'' produces a WD40 repeat. The
UVR8 UV-B resistance 8 (UVR8) also known as ultraviolet-B receptor UVR8 is an UV-B – sensing protein found in plants and possibly other sources. * It is responsible for sensing ultraviolet light in the range 280-315 nm and initiating the plant ...
protein detects UV-B light and mediates the response to this DNA-damaging wavelength. ''A. thaliana'' was used extensively in the study of the genetic basis of
phototropism Phototropism is the growth of an organism in response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi. The cells on the plant that are farthest from the light contain a hor ...
,
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
alignment, and
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
l aperture and other blue light-influenced processes. These traits respond to blue light, which is perceived by the phototropin light receptors. ''Arabidopsis'' has also been important in understanding the functions of another blue light receptor, cryptochrome, which is especially important for light entrainment to control the plants'
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., Endogeny (biology), endogeno ...
s. When the onset of darkness is unusually early, ''A. thaliana'' reduces its metabolism of starch by an amount that effectively requires division. Light responses were even found in roots, previously thought to be largely insensitive to light. While the
gravitropic Gravitropism (also known as geotropism) is a coordinated process of differential growth by a plant in response to gravity pulling on it. It also occurs in fungi. Gravity can be either "artificial gravity" or natural gravity. It is a general featu ...
response of ''A. thaliana'' root organs is their predominant tropic response, specimens treated with
mutagen In genetics, a mutagen is a physical or chemical agent that permanently changes nucleic acid, genetic material, usually DNA, in an organism and thus increases the frequency of mutations above the natural background level. As many mutations can ca ...
s and selected for the absence of gravitropic action showed negative phototropic response to blue or white light, and positive response to red light, indicating that the roots also show positive phototropism. In 2000, Dr. Janet Braam of Rice University genetically engineered ''A. thaliana'' to glow in the dark when touched. The effect was visible to ultrasensitive cameras. Multiple efforts, including the
Glowing Plant project The Glowing Plant project was the first crowdfunding campaign for a synthetic biology application. The project was started by the Sunnyvale-based hackerspace Biocurious as part of the DIYbio philosophy. According to the project's goals, funds we ...
, have sought to use ''A. thaliana'' to increase plant luminescence intensity towards commercially viable levels.


On the Moon

On January 2, 2019, China's
Chang'e-4 Chang'e 4 (; ) is a robotic spacecraft mission, part of the second phase of the Chinese Lunar Exploration Program. China achieved humanity's first soft landing on the far side of the Moon, on 3 January 2019. A communication relay satellit ...
lander brought ''A. thaliana'' to the moon. A small
microcosm Microcosm or macrocosm, also spelled mikrokosmos or makrokosmos, may refer to: Philosophy * Microcosm–macrocosm analogy, the view according to which there is a structural similarity between the human being and the cosmos Music * Macrocosm (alb ...
'tin' in the lander contained ''A. thaliana'', seeds of potatoes, and
silkworm The domestic silk moth (''Bombyx mori''), is an insect from the moth family Bombycidae. It is the closest relative of ''Bombyx mandarina'', the wild silk moth. The silkworm is the larva or caterpillar of a silk moth. It is an economically imp ...
eggs. As plants would support the silkworms with oxygen, and the silkworms would in turn provide the plants with necessary carbon dioxide and nutrients through their waste, researchers will evaluate whether plants successfully perform photosynthesis, and grow and bloom in the lunar environment.


Secondary metabolites

is an ''Arabidopsis'' root triterpene. Potter ''et al.'', 2018 finds
synthesis Synthesis or synthesize may refer to: Science Chemistry and biochemistry *Chemical synthesis, the execution of chemical reactions to form a more complex molecule from chemical precursors ** Organic synthesis, the chemical synthesis of organ ...
is induced by a combination of at least 2 facts, cell-specific transcription factors (TFs) and the accessibility of the chromatin.


Plant–pathogen interactions

Understanding how plants achieve resistance is important to protect the world's food production, and the agriculture industry. Many model systems have been developed to better understand interactions between plants and bacterial, fungal, oomycete,
viral Viral means "relating to viruses" (small infectious agents). Viral may also refer to: Viral behavior, or virality Memetic behavior likened that of a virus, for example: * Viral marketing, the use of existing social networks to spread a marke ...
, and
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
pathogens. ''A. thaliana'' has been a powerful tool for the study of the subdiscipline of plant pathology, that is, the interaction between plants and disease-causing pathogens. The use of ''A. thaliana'' has led to many breakthroughs in the advancement of knowledge of how plants manifest plant disease resistance. The reason most plants are resistant to most pathogens is through nonhost resistance - not all pathogens will infect all plants. An example where ''A. thaliana'' was used to determine the genes responsible for nonhost resistance is '' Blumeria graminis'', the causal agent of powdery mildew of grasses. ''A. thaliana'' mutants were developed using the
mutagen In genetics, a mutagen is a physical or chemical agent that permanently changes nucleic acid, genetic material, usually DNA, in an organism and thus increases the frequency of mutations above the natural background level. As many mutations can ca ...
ethyl methanesulfonate and screened to identify mutants with increased infection by ''B. graminis''. The mutants with higher infection rates are referred to as'' PEN ''mutants due to the ability of ''B. graminis'' to penetrate ''A. thaliana'' to begin the disease process. The ''PEN'' genes were later mapped to identify the genes responsible for nonhost resistance to ''B. graminis''. In general, when a plant is exposed to a pathogen, or nonpathogenic microbe, an initial response, known as PAMP-triggered immunity (PTI), occurs because the plant detects conserved motifs known as pathogen-associated molecular patterns (PAMPs). These PAMPs are detected by specialized receptors in the host known as pattern recognition receptors (PRRs) on the plant cell surface. The best-characterized PRR in ''A. thaliana'' is FLS2 (Flagellin-Sensing2), which recognizes bacterial flagellin, a specialized organelle used by microorganisms for the purpose of motility, as well as the ligand flg22, which comprises the 22 amino acids recognized by FLS2. Discovery of FLS2 was facilitated by the identification of an ''A. thaliana'' ecotype, Ws-0, that was unable to detect flg22, leading to the identification of the gene encoding FLS2. FLS2 shows striking similarity to rice XA21, the first PRR isolated in 1995. Both flagellin and UV-C act similarly to increase homologous recombination in ''A. thaliana'', as demonstrated by Molinier et al. 2006. Beyond this
somatic Somatic may refer to: * Somatic (biology), referring to the cells of the body in contrast to the germ line cells ** Somatic cell, a non-gametic cell in a multicellular organism * Somatic nervous system, the portion of the vertebrate nervous sys ...
effect, they found this to extend to subsequent generations of the plant. A second PRR, EF-Tu receptor (EFR), identified in ''A. thaliana'', recognizes the bacterial EF-Tu protein, the prokaryotic elongation factor used in
protein synthesis Protein biosynthesis (or protein synthesis) is a core biological process, occurring inside Cell (biology), cells, homeostasis, balancing the loss of cellular proteins (via Proteolysis, degradation or Protein targeting, export) through the product ...
, as well as the laboratory-used ligand elf18. Using ''Agrobacterium''-mediated transformation, a technique that takes advantage of the natural process by which ''Agrobacterium'' transfers genes into host plants, the EFR gene was transformed into '' Nicotiana benthamiana'', tobacco plant that does not recognize EF-Tu, thereby permitting recognition of bacterial EF-Tu thereby confirming EFR as the receptor of EF-Tu. Both FLS2 and EFR use similar
signal transduction Signal transduction is the process by which a chemical or physical signal is transmitted through a cell as a series of molecular events, most commonly protein phosphorylation catalyzed by protein kinases, which ultimately results in a cellula ...
pathways to initiate PTI. ''A. thaliana'' has been instrumental in dissecting these pathways to better understand the regulation of immune responses, the most notable one being the
mitogen-activated protein kinase A mitogen-activated protein kinase (MAPK or MAP kinase) is a type of protein kinase that is specific to the amino acids serine and threonine (i.e., a serine/threonine-specific protein kinase). MAPKs are involved in directing cellular responses to ...
(MAP kinase) cascade. Downstream responses of PTI include callose deposition, the oxidative burst, and transcription of defense-related genes. PTI is able to combat pathogens in a nonspecific manner. A stronger and more specific response in plants is that of effector-triggered immunity (ETI), which is dependent upon the recognition of pathogen effectors, proteins secreted by the pathogen that alter functions in the host, by plant resistance genes (R-genes), often described as a gene-for-gene relationship. This recognition may occur directly or indirectly via a guardee protein in a hypothesis known as the guard hypothesis. The first R-gene cloned in ''A. thaliana'' was ''RPS2'' (resistance to ''Pseudomonas syringae'' 2), which is responsible for recognition of the effector avrRpt2. The bacterial effector avrRpt2 is delivered into ''A. thaliana'' via the Type III secretion system of ''P. syringae'' pv. ''tomato'' strain DC3000. Recognition of avrRpt2 by RPS2 occurs via the guardee protein RIN4, which is cleaved. Recognition of a pathogen effector leads to a dramatic immune response known as the hypersensitive response, in which the infected plant cells undergo cell death to prevent the spread of the pathogen. Systemic acquired resistance (SAR) is another example of resistance that is better understood in plants because of research done in ''A. thaliana''. Benzothiadiazol (BTH), a
salicylic acid Salicylic acid is an organic compound with the formula HOC6H4CO2H. A colorless, bitter-tasting solid, it is a precursor to and a metabolite of aspirin (acetylsalicylic acid). It is a plant hormone, and has been listed by the EPA Toxic Substance ...
(SA) analog, has been used historically as an antifungal compound in crop plants. BTH, as well as SA, has been shown to induce SAR in plants. The initiation of the SAR pathway was first demonstrated in ''A. thaliana'' in which increased SA levels are recognized by nonexpresser of PR genes 1 (''NPR1'') due to redox change in the cytosol, resulting in the reduction of ''NPR1. NPR1'', which usually exists in a multiplex (oligomeric) state, becomes monomeric (a single unit) upon reduction. When NPR1 becomes monomeric, it translocates to the nucleus, where it interacts with many TGA transcription factors, and is able to induce pathogen-related genes such as ''PR1''. Another example of SAR would be the research done with transgenic tobacco plants, which express bacterial salicylate hydroxylase, nahG gene, requires the accumulation of SA for its expression Although not directly immunological, intracellular transport affects susceptibility by incorporating - or being tricked into incorporating - pathogen particles. For example, the '' Dynamin-related protein 2b/
drp2b DRP may refer to: Political parties * Democratic Republican Party (South Korea) * Deutsche Rechtspartei, a right-wing German party * Dhivehi Rayyithunge Party, a right-wing Maldivian party Other uses * Dells Raceway Park, Wisconsin, US * Distribut ...
'' gene helps to move invaginated material into cells, with some mutants increasing '' PstDC3000'' virulence even further.


Evolutionary aspect of plant-pathogen resistance

Plants are affected by multiple pathogens throughout their lifetimes. In response to the presence of pathogens, plants have evolved receptors on their cell surfaces to detect and respond to pathogens. ''Arabidopsis thaliana'' is a model organism used to determine specific defense mechanisms of plant-pathogen resistance. These plants have special receptors on their cell surfaces that allow for detection of pathogens and initiate mechanisms to inhibit pathogen growth. They contain two receptors, FLS2 (bacterial flagellin receptor) and EF-Tu (bacterial EF-Tu protein), which use signal transduction pathways to initiate the disease response pathway. The pathway leads to the recognition of the pathogen causing the infected cells to undergo cell death to stop the spread of the pathogen. Plants with FLS2 and EF-Tu receptors have shown to have increased fitness in the population. This has led to the belief that plant-pathogen resistance is an evolutionary mechanism that has built up over generations to respond to dynamic environments, such as increased predation and extreme temperatures. ''A. thaliana'' has also been used to study SAR. This pathway uses benzothiadiazol, a chemical inducer, to induce transcription factors, mRNA, of SAR genes. This accumulation of transcription factors leads to inhibition of pathogen-related genes. Plant-pathogen interactions are important for an understanding of how plants have evolved to combat different types of pathogens that may affect them. Variation in resistance of plants across populations is due to variation in environmental factors. Plants that have evolved resistance, whether it be the general variation or the SAR variation, have been able to live longer and hold off necrosis of their tissue (premature death of cells), which leads to better adaptation and fitness for populations that are in rapidly changing environments. In the future, comparisons of the pathosystems of wild populations + their coevolved pathogens with wild-wild hybrids of known parentage may reveal new mechanisms of balancing selection. In life history theory we may find that ''A. thaliana'' maintains certain alleles due to
pleitropy Pleiotropy (from Greek , 'more', and , 'way') occurs when one gene influences two or more seemingly unrelated phenotypic traits. Such a gene that exhibits multiple phenotypic expression is called a pleiotropic gene. Mutation in a pleiotropic g ...
between plant-pathogen effects and other traits, as in livestock. Research in ''A. thaliana'' suggests that the immunity regulator protein family EDS1 in general co-evolved with the CC family of nucleotide-bindingleucine-rich-repeat-receptors (NLRs). Xiao et al. 2005 have shown that the powdery mildew immunity mediated by ''A. thaliana''s RPW8 (which has a CC
domain Domain may refer to: Mathematics *Domain of a function, the set of input values for which the (total) function is defined **Domain of definition of a partial function **Natural domain of a partial function **Domain of holomorphy of a function * Do ...
) is dependent on two members of this family: '' EDS1'' itself and '' PAD4''. ''
RESISTANCE TO PSEUDOMONAS SYRINGAE 5 Resistance may refer to: Arts, entertainment, and media Comics * Either of two similarly named but otherwise unrelated comic book series, both published by Wildstorm: ** Resistance (comics), ''Resistance'' (comics), based on the video game of th ...
/RPS5'' is a disease resistance protein which guards ''
AvrPphB SUSCEPTIBLE 1 ''AvrPphB SUSCEPTIBLE 1/PBS1'' is a protein kinase acting upon serine and threonine. . . . It is a receptor-like cytoplasmic kinase, of Subfamily VII. It is the guardee of ''RESISTANCE TO PSEUDOMONAS SYRINGAE 5/RPS5''. PBS1 is cleaved by an effecto ...
/PBS1''. ''PBS1'', as the name would suggest, is the target of '' AvrPphB'', an effector produced by ''Pseudomonas syringae'' pv. ''phaseolicola''.


Other research

Ongoing research on ''A. thaliana'' is being performed on the International Space Station by the
European Space Agency , owners = , headquarters = Paris, Île-de-France, France , coordinates = , spaceport = Guiana Space Centre , seal = File:ESA emblem seal.png , seal_size = 130px , image = Views in the Main Control Room (1205 ...
. The goals are to study the growth and reproduction of plants from seed to seed in
microgravity The term micro-g environment (also μg, often referred to by the term microgravity) is more or less synonymous with the terms ''weightlessness'' and ''zero-g'', but emphasising that g-forces are never exactly zero—just very small (on the I ...
. Plant-on-a-chip devices in which ''A. thaliana'' tissues can be cultured in semi-''in vitro'' conditions have been described. Use of these devices may aid understanding of pollen-tube guidance and the mechanism of sexual reproduction in ''A. thaliana.'' Researchers at the University of Florida were able to grow the plant in lunar soil originating from the Sea of Tranquillity.


Self-pollination

''A. thaliana'' is a predominantly self-pollinating plant with an outcrossing rate estimated at less than 0.3%. An analysis of the genome-wide pattern of linkage disequilibrium suggested that self-pollination evolved roughly a million years ago or more. Meioses that lead to self-pollination are unlikely to produce significant beneficial genetic variability. However, these meioses can provide the adaptive benefit of recombinational repair of DNA damages during formation of germ cells at each generation. Such a benefit may have been sufficient to allow the long-term persistence of meioses even when followed by self-fertilization. A physical mechanism for self-pollination in ''A. thaliana'' is through pre-anthesis autogamy, such that fertilisation takes place largely before flower opening.


Databases and other resources

* TAIR and NASC: curated sources for diverse genetic and molecular biology information, links to gene expression databases etc. * Arabidopsis Biological Resource Center (seed and DNA stocks) * Nottingham Arabidopsis Stock Centre (seed and DNA stocks) *
Artade ARTADE (ARabidopsis Tiling Array-based Detection of Exons) is a database for the annotation of genome-wide tiling-array data in Arabidopsis See also * DNA microarray A DNA microarray (also commonly known as DNA chip or biochip) is a collecti ...
database


See also

* ''A. thaliana'' responses to salinity * BZIP intron plant * The Thaliana Bridge, installed in 2021 at
Harlow Carr Harlow is a large town and local government district located in the west of Essex, England. Founded as a new town, it is situated on the border with Hertfordshire and London, Harlow occupies a large area of land on the south bank of the upper ...
was inspired by the work of the botanical scientist Rachel Leech and represents the sequence of an ''Arabidopsis thaliana'' chromosome.


References


External links

*
Arabidopsis transcriptional regulatory mapThe Arabidopsis Information Resource (TAIR)What Makes Plants Grow? The Arabidopsis genome knows
Featured article in Genome News Network
The Arabidopsis book
- A comprehensive review published yearly related to research in ''Arabidopsis''
A. thaliana protein abundanceThe Arabidopsis Information Portal (Araport)
{{Use dmy dates, date=January 2015 Flora of Europe Flora of Asia Flora of Africa Flora of Lebanon Plant models Plants described in 1753 Taxa named by Carl Linnaeus Space-flown life Edible plants Plant cognition thaliana