Woese's dogma is a principle of evolutionary biology first put forth by biophysicist Carl Woese in 1977. It states that the evolution of ribosomal RNA was a necessary precursor to the evolution of modern life forms. [1] This led to the advancement of the phylogenetic tree of life consisting of three domains rather than the previously accepted two. [2] While the existence of Eukarya and Prokarya were already accepted, Woese was responsible for the distinction between Bacteria and Archaea. [3] Despite initial criticism and controversy surrounding his claims, Woese's three domain system, based on his work regarding the role of rRNA in the evolution of modern life, has become widely accepted. [4]
Evidence for Woese's dogma is well established through comparisons of RNA homology. Modern research allows more liberal use of RNA sequencing, allowing for a better comparative analysis between distant RNA. When analyzing multiple strains of E. coli, Root-Bernstein et al. have compared tRNA encodings found within rRNA with tRNA found in E. coli to see if the secondary structure was the same as more “modern” tRNA present in E. coli. [5] Comparisons between the tRNA encodings found in the rRNAs and mRNAs of the control sequences found that “sortings” for these sequences were extremely similar, and comparisons of translated protein structure indicated that homology was likely. [5] Additionally, sequences homologous to all tRNAs necessary for translation were present in 16s and 23s rRNAs, and synthetases to load these tRNAs were also found, indicating that many of the functions of transcription and translation present in more modern life exist in rRNA, if vestigially. [5]
When comparing homologies of rRNA structures, it is necessary to analyze substructures. This is because models that study RNA structure on the whole do not currently exist. [6] Generally, phylogenies of rRNA subunits are created to understand each component, and how they function and evolve. Through phylogenies created that depict rRNA structural elements that are present in all three domains of life, the oldest structural components can be determined through relative dating. [7] These phylogenies were used in a study by Harish et al., to show that a helical stem labeled h44 in small subunit rRNA can be described as the oldest structural component of rRNA, which holds particular significance, as this structure responsible for linking processes in the small subunit, which is responsible for decoding, with the large subunit, which is responsible for the formation of peptide bonds and the releasing of elongation factors. [7] This essentially shows that the functional origin of the ribosome, responsible for protein synthesis, is common in all modern life throughout each of the three domains.
Evidence has also been obtained in studying eukaryotic organelles, such as the chloroplast. Zablen et al.’s phylogenetic analysis conducted electrophoresis on chloroplast ribosomal RNA, specifically on the 16S rRNA of Euglena gracilis . [8] In conducting this experiment, researchers compared the electrophoretic fingerprint of this RNA to other chloroplasts and prokarya. In comparing these results, it was found that generally, these chloroplasts show a close genomic relationship, while a more distant one is seen for algae, and subsequently prokaryotic organisms. [8] This experiment shows that the rRNA of distantly related organisms has a similar origin of that in eukaryotic organelles, supporting the idea that the evolution of rRNA was a necessary precursor of modern life.
One of the reasons that Woese's Dogma holds significance is because of the potential that RNA was the first primordial self-replicating molecule (see: RNA World), meaning it would be key in the progression of modern life. [9] In particular, it has been proposed that ribosomes exist as a missing link in prebiotic evolution, with rRNA being a vestige of an ancient genome. [5] Some evidence exists for the proposal that rRNA functioned in the past to encode proteins that are key to ribosome function. [10] One notable example is the fact that rRNA proteins are commonly known to bind with their own mRNA. [10] In addition, some ribosomal proteins not only regulate their own expression, but the expression of other proteins as well. [11] These are both indications of self-replication, and indicate the possibility that the mRNA that encodes ribosomal proteins evolved from rRNA. [10]
RNA existing as a primordial self replicating entity is an idea that faces criticism. The idea of rRNA in particular being sufficient on its own to explain the progression of modern life struggles due to the fact that it lacks certain key pieces of evidence. In particular RNA cannot be shown to be prebiotic, as there is no way for the nucleotides or nucleosides that compose it to be non-enzymatically replicated. [12] Additionally, other criticisms exist, such as the fact that RNA is not stable enough to have arisen prebiotically, and that it is too complex to have arisen prebiotically. [13] This has led to the development of other hypotheses, such as 'proteins first', which states that proteins arose prior to RNA, or coevolved with RNA. [13] This has also led to the proposal of other primordial molecules that may have developed into RNA and DNA, such as peptide nucleic acids, which also show evidence of self replication. [14] Despite the fact that criticisms might exist on the primordial or prebiotic nature of rRNA, these criticisms are not aimed at Woese's Dogma on the whole, as Woese's Dogma only claims that the evolution of rRNA was a necessary precursor to modern life, not that rRNA arose prebiotically. [1]
Ribonucleic acid (RNA) is a polymeric molecule that is essential for most biological functions, either by performing the function itself or by forming a template for the production of proteins. RNA and deoxyribonucleic acid (DNA) are nucleic acids. The nucleic acids constitute one of the four major macromolecules essential for all known forms of life. RNA is assembled as a chain of nucleotides. Cellular organisms use messenger RNA (mRNA) to convey genetic information that directs synthesis of specific proteins. Many viruses encode their genetic information using an RNA genome.
The RNA world is a hypothetical stage in the evolutionary history of life on Earth in which self-replicating RNA molecules proliferated before the evolution of DNA and proteins. The term also refers to the hypothesis that posits the existence of this stage.
Ribosomes are macromolecular machines, found within all cells, that perform biological protein synthesis. Ribosomes link amino acids together in the order specified by the codons of messenger RNA molecules to form polypeptide chains. Ribosomes consist of two major components: the small and large ribosomal subunits. Each subunit consists of one or more ribosomal RNA molecules and many ribosomal proteins. The ribosomes and associated molecules are also known as the translational apparatus.
The central dogma of molecular biology deals with the flow of genetic information within a biological system. It is often stated as "DNA makes RNA, and RNA makes protein", although this is not its original meaning. It was first stated by Francis Crick in 1957, then published in 1958:
The Central Dogma. This states that once "information" has passed into protein it cannot get out again. In more detail, the transfer of information from nucleic acid to nucleic acid, or from nucleic acid to protein may be possible, but transfer from protein to protein, or from protein to nucleic acid is impossible. Information here means the precise determination of sequence, either of bases in the nucleic acid or of amino acid residues in the protein.
In molecular biology, RNA polymerase, or more specifically DNA-directed/dependent RNA polymerase (DdRP), is an enzyme that catalyzes the chemical reactions that synthesize RNA from a DNA template.
Ribozymes are RNA molecules that have the ability to catalyze specific biochemical reactions, including RNA splicing in gene expression, similar to the action of protein enzymes. The 1982 discovery of ribozymes demonstrated that RNA can be both genetic material and a biological catalyst, and contributed to the RNA world hypothesis, which suggests that RNA may have been important in the evolution of prebiotic self-replicating systems.
Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal DNA (rDNA) and then bound to ribosomal proteins to form small and large ribosome subunits. rRNA is the physical and mechanical factor of the ribosome that forces transfer RNA (tRNA) and messenger RNA (mRNA) to process and translate the latter into proteins. Ribosomal RNA is the predominant form of RNA found in most cells; it makes up about 80% of cellular RNA despite never being translated into proteins itself. Ribosomes are composed of approximately 60% rRNA and 40% ribosomal proteins, though this ratio differs between prokaryotes and eukaryotes.
Bacterial translation is the process by which messenger RNA is translated into proteins in bacteria.
A nuclear gene is a gene that has its DNA nucleotide sequence physically situated within the cell nucleus of a eukaryotic organism. This term is employed to differentiate nuclear genes, which are located in the cell nucleus, from genes that are found in mitochondria or chloroplasts. The vast majority of genes in eukaryotes are nuclear.
A ribosomal protein is any of the proteins that, in conjunction with rRNA, make up the ribosomal subunits involved in the cellular process of translation. E. coli, other bacteria and Archaea have a 30S small subunit and a 50S large subunit, whereas humans and yeasts have a 40S small subunit and a 60S large subunit. Equivalent subunits are frequently numbered differently between bacteria, Archaea, yeasts and humans.
The 5S ribosomal RNA is an approximately 120 nucleotide-long ribosomal RNA molecule with a mass of 40 kDa. It is a structural and functional component of the large subunit of the ribosome in all domains of life, with the exception of mitochondrial ribosomes of fungi and animals. The designation 5S refers to the molecule's sedimentation coefficient in an ultracentrifuge, which is measured in Svedberg units (S).
The 23S rRNA is a 2,904 nucleotide long component of the large subunit (50S) of the bacterial/archean ribosome and makes up the peptidyl transferase center (PTC). The 23S rRNA is divided into six secondary structural domains titled I-VI, with the corresponding 5S rRNA being considered domain VII. The ribosomal peptidyl transferase activity resides in domain V of this rRNA, which is also the most common binding site for antibiotics that inhibit translation, making it a target for ribosomal engineering. A well-known member of this antibiotic class, chloramphenicol, acts by inhibiting peptide bond formation, with recent 3D-structural studies showing two different binding sites depending on the species of ribosome. Numerous mutations in domains of the 23S rRNA with Peptidyl transferase activity have resulted in antibiotic resistance. 23S rRNA genes typically have higher sequence variations, including insertions and/or deletions, compared to other rRNAs.
28S ribosomal protein S24, mitochondrial is a protein that in humans is encoded by the MRPS24 gene.
EF-G is a prokaryotic elongation factor involved in mRNA translation. As a GTPase, EF-G catalyzes the movement (translocation) of transfer RNA (tRNA) and messenger RNA (mRNA) through the ribosome.
39S ribosomal protein L10, mitochondrial is a protein that in humans is encoded by the MRPL10 gene.
Evolution of cells refers to the evolutionary origin and subsequent evolutionary development of cells. Cells first emerged at least 3.8 billion years ago approximately 750 million years after Earth was formed.
Chloroplast DNA (cpDNA) is the DNA located in chloroplasts, which are photosynthetic organelles located within the cells of some eukaryotic organisms. Chloroplasts, like other types of plastid, contain a genome separate from that in the cell nucleus. The existence of chloroplast DNA was identified biochemically in 1959, and confirmed by electron microscopy in 1962. The discoveries that the chloroplast contains ribosomes and performs protein synthesis revealed that the chloroplast is genetically semi-autonomous. The first complete chloroplast genome sequences were published in 1986, Nicotiana tabacum (tobacco) by Sugiura and colleagues and Marchantia polymorpha (liverwort) by Ozeki et al. Since then, a great number of chloroplast DNAs from various species have been sequenced.
Numerous key discoveries in biology have emerged from studies of RNA, including seminal work in the fields of biochemistry, genetics, microbiology, molecular biology, molecular evolution, and structural biology. As of 2010, 30 scientists have been awarded Nobel Prizes for experimental work that includes studies of RNA. Specific discoveries of high biological significance are discussed in this article.
Small subunit ribosomal ribonucleic acid is the smaller of the two major RNA components of the ribosome. Associated with a number of ribosomal proteins, the SSU rRNA forms the small subunit of the ribosome. It is encoded by SSU-rDNA.
The first universal common ancestor (FUCA) is a proposed non-cellular entity that was the earliest organism with a genetic code capable of biological translation of RNA molecules into peptides to produce proteins. Its descendents include the last universal common ancestor (LUCA) and every modern cell. FUCA would also be the ancestor of ancient sister lineages of LUCA, none of which have modern descendants, but which are thought to have horizontally transferred some of their genes into the genome of early descendants of LUCA.