The MADS box is a conserved sequence motif. The genes which contain this motif are called the MADS-box gene family. [1] The MADS box encodes the DNA-binding MADS domain. The MADS domain binds to DNA sequences of high similarity to the motif CC[A/T]6GG termed the CArG-box. [2] MADS-domain proteins are generally transcription factors. [2] [3] The length of the MADS-box reported by various researchers varies somewhat, but typical lengths are in the range of 168 to 180 base pairs, i.e. the encoded MADS domain has a length of 56 to 60 amino acids. [4] [5] [6] [7] There is evidence that the MADS domain evolved from a sequence stretch of a type II topoisomerase in a common ancestor of all extant eukaryotes. [8]
The first MADS-box gene to be identified was ARG80 from budding yeast, Saccharomyces cerevisiae , [9] but was at that time not recognized as a member of a large gene family. The MADS-box gene family got its name later as an acronym referring to the four founding members, [1] ignoring ARG80:
In A. thaliana, A. majus, and Zea mays this motif is involved in floral development. Early study in these model angiosperms was the beginning of research into the molecular evolution of floral structure in general, as well as their role in nonflowering plants. [11]
MADS-box genes have been detected in nearly all eukaryotes studied. [8] While the genomes of animals and fungi generally possess only around one to five MADS-box genes, genomes of flowering plants have around 100 MADS-box genes. [12] [13] Two types of MADS-domain proteins are distinguished; the SRF-like or Type I MADS-domain proteins and the MEF2-like (after MYOCYTE-ENHANCER-FACTOR2) or Type II MADS-domain proteins. [8] [13] SRF-like MADS-domain proteins in animals and fungi have a second conserved domain, the SAM (SRF, ARG80, MCM1) domain. [14] MEF2-like MADS-domain proteins in animals and fungi have the MEF2 domain as a second conserved domain. [14] In plants, the MEF2-like MADS-domain proteins are also termed MIKC-type proteins referring to their conserved domain structure, where the MADS (M) domain is followed by an Intervening (I), a Keratin-like (K) and a C-terminal domain. [12] In plants, MADS-domain protein form tetramers and this is thought to be central for their function. [15] [16] The structure of the tetramerisation domain of the MADS-domain protein SEPALLATA3 was solved illustrating the structural basis for tetramer formation [17]
A geneticist intensely investigating MADS-box genes is Günter Theißen at the University of Jena. For example, he and his coworkers have used these genes to show that the order Gnetales is more closely related to the conifers than to the flowering plants. [18]
MADS-box is under-studied in wheat as of 2021 [update] . [19]
In Zea mays the mutant Tunicate1 produces pod corn. Tunicate1 is a mutant of Z. mays MADS19 (ZMM19), in the SHORT VEGETATIVE PHASE gene family. ZMM19 can be ectopically expressed. [19]
Such ectopic expression of ZMM19 in A. thaliana enlarges sepals, suggesting conservation. [19]
MADS-box genes have a variety of functions. In animals, MADS-box genes are involved in muscle development and cell proliferation and differentiation. [14] Functions in fungi range from pheromone response to arginine metabolism. [14]
In plants, MADS-box genes are involved in controlling all major aspects of development, including male and female gametophyte development, embryo and seed development, as well as root, flower and fruit development. [12] [13]
Some MADS-box genes of flowering plants have homeotic functions like the HOX genes of animals. [1] The floral homeotic MADS-box genes (such as AGAMOUS and DEFICIENS) participate in the determination of floral organ identity according to the ABC model of flower development. [20]
Another function of MADS-box genes is flowering time determination. In Arabidopsis thaliana the MADS box genes SOC1 [21] and Flowering Locus C [22] (FLC) have been shown to have an important role in the integration of molecular flowering time pathways. These genes are essential for the correct timing of flowering , and help to ensure that fertilization occurs at the time of maximal reproductive potential.
The MADS box protein structure is characterized by four domains. At the N terminal end is the highly conserved MADS DNA binding domain. [23] Next to the MADS domain is the moderately conserved Intervening (I) and Keratin-like (K) domains, which are involved in specific protein-protein interactions. [23] The carboxyl terminal (C) domain is highly variable and is involved in transcriptional activation and assemblage of heterodimers and multimeric protein complexes. [24]
Arabidopsis thaliana, the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally considered a weed.
In cell biology, the meristem is a type of tissue found in plants. It consists of undifferentiated cells capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells continue to divide until they become differentiated and lose the ability to divide.
In evolutionary developmental biology, homeosis is the transformation of one organ into another, arising from mutation in or misexpression of certain developmentally critical genes, specifically homeotic genes. In animals, these developmental genes specifically control the development of organs on their anteroposterior axis. In plants, however, the developmental genes affected by homeosis may control anything from the development of a stamen or petals to the development of chlorophyll. Homeosis may be caused by mutations in Hox genes, found in animals, or others such as the MADS-box family in plants. Homeosis is a characteristic that has helped insects become as successful and diverse as they are.
Florigens are proteins capable of inducing flowering time in angiosperms. The prototypical florigen is encoded by the FT gene and its orthologs in Arabidopsis and other plants. Florigens are produced in the leaves, and act in the shoot apical meristem of buds and growing tips.
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 are three physiological developments that must occur in order for this to take place: firstly, the plant must pass from sexual immaturity into a sexually mature state ; secondly, the transformation of the apical meristem's function from a vegetative meristem into a floral meristem or inflorescence; and finally the growth of the flower's individual organs. The latter phase has been modelled using the ABC model, which aims to describe the biological basis of the process from the perspective of molecular and developmental genetics.
Polycomb-group proteins are a family of protein complexes first discovered in fruit flies that can remodel chromatin such that epigenetic silencing of genes takes place. Polycomb-group proteins are well known for silencing Hox genes through modulation of chromatin structure during embryonic development in fruit flies. They derive their name from the fact that the first sign of a decrease in PcG function is often a homeotic transformation of posterior legs towards anterior legs, which have a characteristic comb-like set of bristles.
In the field of molecular biology, myocyte enhancer factor-2 (Mef2) proteins are a family of transcription factors which through control of gene expression are important regulators of cellular differentiation and consequently play a critical role in embryonic development. In adult organisms, Mef2 proteins mediate the stress response in some tissues. Mef2 proteins contain both MADS-box and Mef2 DNA-binding domains.
Apetala 2(AP2) is a gene and a member of a large family of transcription factors, the AP2/EREBP family. In Arabidopsis thaliana AP2 plays a role in the ABC model of flower development. It was originally thought that this family of proteins was plant-specific; however, recent studies have shown that apicomplexans, including the causative agent of malaria, Plasmodium falciparum encode a related set of transcription factors, called the ApiAP2 family.
Evolutionary developmental biology (evo-devo) is the study of developmental programs and patterns from an evolutionary perspective. It seeks to understand the various influences shaping the form and nature of life on the planet. Evo-devo arose as a separate branch of science rather recently. An early sign of this occurred in 1999.
Myocyte-specific enhancer factor 2C also known as MADS box transcription enhancer factor 2, polypeptide C is a protein that in humans is encoded by the MEF2C gene. MEF2C is a transcription factor in the Mef2 family.
Myocyte-specific enhancer factor 2A is a protein that in humans is encoded by the MEF2A gene. MEF2A is a transcription factor in the Mef2 family. In humans it is located on chromosome 15q26. Certain mutations in MEF2A cause an autosomal dominant form of coronary artery disease and myocardial infarction.
GAI or Gibberellic-Acid Insensitive is a gene in Arabidopsis thaliana which is involved in regulation of plant growth. GAI represses the pathway of gibberellin-sensitive plant growth. It does this by way of its conserved DELLA motif.
Ethylene-responsive element binding protein(EREBP) is a homeobox gene from Arabidopsis thaliana and other plants which encodes a transcription factor. EREBP is responsible in part for mediating the response in plants to the plant hormone ethylene.
Agamous (AG) is a homeotic gene and MADS-box transcription factor from Arabidopsis thaliana. The TAIR AGI number is AT4G18960.
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 is expressed, and flowering becomes possible. FLC is extensively regulated through epigenetic modifications and transcriptional control.
Homeotic genes are genes which regulate the development of anatomical structures in various organisms such as echinoderms, insects, mammals, and plants. Homeotic genes often encode transcription factor proteins, and these proteins affect development by regulating downstream gene networks involved in body patterning.
LEAFY is a plant gene that causes groups of undifferentiated cells called meristems to develop into flowers instead of leaves with associated shoots.
Detlef Weigel is a German American scientist working at the interface of developmental and evolutionary biology.
The WRKY domain is found in the WRKY transcription factor family, a class of transcription factors. The WRKY domain is found almost exclusively in plants although WRKY genes appear present in some diplomonads, social amoebae and other amoebozoa, and fungi incertae sedis. They appear absent in other non-plant species. WRKY transcription factors have been a significant area of plant research for the past 20 years. The WRKY DNA-binding domain recognizes the W-box (T)TGAC(C/T) cis-regulatory element.
EARLY FLOWERING 3 (ELF3) is a plant-specific gene that encodes the hydroxyproline-rich glycoprotein and is required for the function of the circadian clock. ELF3 is one of the three components that make up the Evening Complex (EC) within the plant circadian clock, in which all three components reach peak gene expression and protein levels at dusk. ELF3 serves as a scaffold to bind EARLY FLOWERING 4 (ELF4) and LUX ARRHYTHMO (LUX), two other components of the EC, and functions to control photoperiod sensitivity in plants. ELF3 also plays an important role in temperature and light input within plants for circadian clock entrainment. Additionally, it plays roles in light and temperature signaling that are independent from its role in the EC.
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