Reductive evolution is the process by which microorganisms remove genes from their genome. It can occur when bacteria found in a free-living state enter a restrictive state (either as endosymbionts or parasites) or are completely absorbed by another organism becoming intracellular (symbiogenesis). The bacteria will adapt to survive and thrive in the restrictive state by altering and reducing its genome to get rid of the newly redundant pathways that are provided by the host. [1] In an endosymbiont or symbiogenesis relationship where both the guest and host benefit, the host can also undergo reductive evolution to eliminate pathways that are more efficiently provided for by the guest. [2]
Endosymbiont or parasitic microorganisms such as Rickettsia prowazekii , Chlorella in Paramecium , Buchnera aphidicola in aphids, and Wolbachia bacteria in Wuchereria bancrofti have all been studied and fully sequenced which is why they are used as examples of reductive evolution. Sometimes bacteria will eliminate genes from their genome, this is called reductive evolution. Reductive genes can be nonessential to the organism and makes it so the bacteria can reproduce more efficiently. [1]
Another example of this would be the black queen hypothesis, where bacteria rely on extracellular metabolites, produced by symbiotic bacteria in their environment. The bacteria become dependent on one another by reducing, getting rid of the genes responsible for producing their own metabolites. It can also be a from obligate intracellular organisms that reduce their genomes and become dependent on the host to produce metabolites for the organism to use. [3]
Reductive evolution [4] is the basis behind the Endosymbiotic Theory, which states that Eukaryotes absorbed other microorganisms (Eukaryotes and archaea) for their metabolites produced. The absorbed organisms undergo reductive evolution, deleting genes that were deemed nonessential or non-beneficial to the cell in its specific niche in the host. When comparing fossil evidence reductive evolution can be demonstrated. [5]
DNA found in ancient prokaryotic and mitochondria fossils have been found to have higher levels of cytosine and guanine compared to the DNA found in the same organism today. Different segments of the genome found to be unfavorable have possibly been removed over time due to deletions of DNA causing the genome to be reduced. [6] The amount of cytosine and guanine in an organism's genome is a direct correlation to the overall size of that genome. [7]
The genome can become more complex or simplified due to random mutations. [8]
Chlorella is a secondary endosymbiont that lives within Paramecium species and is an example of obligate intracellular reductive evolution. Moranella is a double membrane gram-negative-like bacteria that lives in another endosymbiont, "Candidatus Tremblaya", which itself lives in the mealy bug.
Following reductive evolution, it is suggested that between 180 and 425 million years ago the Rickettsia parasite incident occurred. It has been hypothesized that this event had to have happened later on as the Rickettsia and mitochondria evolved from a common ancestor. With this information, scientists understand that Rickettsia and mitochondria had to have happened at different points in their evolution. Fossils have been used to identify and confirm these endosymbiotic events, but not nearly enough have been found for a good statistical sample size. [6]
Lyn Margulis remarked, "bacterium established a stable residence within the cytoplasm of a primitive eukaryote and supplied the cell with energy in exchange for a protected environment with a ready supply of nutrients." [9] This became the leading theory of endosymbiosis. This was further proved with the finding that mitochondria and chloroplasts had a separate genome from the host genome, but had lost the ability to live outside of the host.
There are many methods to help identify if genes have been deleted, two of which are maximum parsimony (MP) or maximum likelihood (ML) patterns are used to recreate the evolutionary tree of these species and their gene compositions of the ancient forms as well as the gene losses and gained along the tree branches which are then compared to each other. There are limitations, however, mostly due to using different models or adding new information which can skew results. Such as using Dollo Parsimony or Weighted Parsimony.
Maximum parsimony (MP)
Maximum likelihood (ML)
Rickettsia prowazekii is an unrestricted microorganism which has been used to demonstrate genome degradation [10] DNA and genome size is not linked to the complexity of an organism. There are some bacteria that have a lot more DNA than a human. This is not yet understood and is referred to as the C-value Enigma or C-value Paradox. In other words, the vast amount of DNA in a haploid genome doesn't compare to the complexity of an organism and can be very different. Through the process of reductive evolution large sections of the DNA could have been removed, turned off, or phased out by the organism if found to be no longer useful in its desire to survive and grow.
An endosymbiont or endobiont is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living".) Examples are nitrogen-fixing bacteria, which live in the root nodules of legumes, single-cell algae inside reef-building corals and bacterial endosymbionts that provide essential nutrients to insects.
Symbiosis is any type of a close and long-term biological interaction between two biological organisms of different species, termed symbionts, be it mutualistic, commensalistic, or parasitic. In 1879, Heinrich Anton de Bary defined it as "the living together of unlike organisms". The term is sometimes used in the more restricted sense of a mutually beneficial interaction in which both symbionts contribute to each other's support.
Symbiogenesis is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibly other organelles of eukaryotic cells are descended from formerly free-living prokaryotes taken one inside the other in endosymbiosis. Mitochondria appear to be phylogenetically related to Rickettsiales bacteria, while chloroplasts are thought to be related to cyanobacteria.
Rickettsia is a genus of nonmotile, gram-negative, nonspore-forming, highly pleomorphic bacteria that may occur in the forms of cocci, bacilli, or threads. The genus was named after Howard Taylor Ricketts in honor of his pioneering work on tick-borne spotted fever.
Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the evolution of many organisms. HGT is influencing scientific understanding of higher order evolution while more significantly shifting perspectives on bacterial evolution.
Nucleomorphs are small, vestigial eukaryotic nuclei found between the inner and outer pairs of membranes in certain plastids. They are thought to be vestiges of primitive red and green algal nuclei that were engulfed by a larger eukaryote. Because the nucleomorph lies between two sets of membranes, nucleomorphs support the endosymbiotic theory and are evidence that the plastids containing them are complex plastids. Having two sets of membranes indicate that the plastid, a prokaryote, was engulfed by a eukaryote, an alga, which was then engulfed by another eukaryote, the host cell, making the plastid an example of secondary endosymbiosis.
Rickettsia prowazekii is a species of gram-negative, alphaproteobacteria, obligate intracellular parasitic, aerobic bacillus bacteria that is the etiologic agent of epidemic typhus, transmitted in the feces of lice. In North America, the main reservoir for R. prowazekii is the flying squirrel. R. prowazekii is often surrounded by a protein microcapsular layer and slime layer; the natural life cycle of the bacterium generally involves a vertebrate and an invertebrate host, usually an arthropod, typically the human body louse. A form of R. prowazekii that exists in the feces of arthropods remains stably infective for months. R. prowazekii also appears to be the closest free-living relative of mitochondria, based on genome sequencing.
The Rickettsiales, informally called rickettsias, are an order of small Alphaproteobacteria. They are obligate intracellular parasites, and some are notable pathogens, including Rickettsia, which causes a variety of diseases in humans, and Ehrlichia, which causes diseases in livestock. Another genus of well-known Rickettsiales is the Wolbachia, which infect about two-thirds of all arthropods and nearly all filarial nematodes. Genetic studies support the endosymbiotic theory according to which mitochondria and related organelles developed from members of this group.
Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms or less frequently in daltons, or as the total number of nucleotide base pairs, usually in megabases. One picogram is equal to 978 megabases. In diploid organisms, genome size is often used interchangeably with the term C-value.
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.
"Candidatus Carsonella ruddii" is an obligate endosymbiotic Gammaproteobacterium with one of the smallest genomes of any characterised bacteria.
Ehrlichia is a genus of Rickettsiales bacteria that are transmitted to vertebrates by ticks. These bacteria cause the disease ehrlichiosis, which is considered zoonotic, because the main reservoirs for the disease are animals.
A prokaryote is a single-cell organism whose cell lacks a nucleus and other membrane-bound organelles. The word prokaryote comes from the Ancient Greek πρό 'before' and κάρυον 'nut, kernel'. In the two-empire system arising from the work of Édouard Chatton, prokaryotes were classified within the empire Prokaryota. But in the three-domain system, based upon molecular analysis, prokaryotes are divided into two domains: Bacteria and Archaea. Organisms with nuclei are placed in a third domain, Eukaryota.
Bacterial genomes are generally smaller and less variant in size among species when compared with genomes of eukaryotes. Bacterial genomes can range in size anywhere from about 130 kbp to over 14 Mbp. A study that included, but was not limited to, 478 bacterial genomes, concluded that as genome size increases, the number of genes increases at a disproportionately slower rate in eukaryotes than in non-eukaryotes. Thus, the proportion of non-coding DNA goes up with genome size more quickly in non-bacteria than in bacteria. This is consistent with the fact that most eukaryotic nuclear DNA is non-gene coding, while the majority of prokaryotic, viral, and organellar genes are coding. Right now, we have genome sequences from 50 different bacterial phyla and 11 different archaeal phyla. Second-generation sequencing has yielded many draft genomes ; third-generation sequencing might eventually yield a complete genome in a few hours. The genome sequences reveal much diversity in bacteria. Analysis of over 2000 Escherichia coli genomes reveals an E. coli core genome of about 3100 gene families and a total of about 89,000 different gene families. Genome sequences show that parasitic bacteria have 500–1200 genes, free-living bacteria have 1500–7500 genes, and archaea have 1500–2700 genes. A striking discovery by Cole et al. described massive amounts of gene decay when comparing Leprosy bacillus to ancestral bacteria. Studies have since shown that several bacteria have smaller genome sizes than their ancestors did. Over the years, researchers have proposed several theories to explain the general trend of bacterial genome decay and the relatively small size of bacterial genomes. Compelling evidence indicates that the apparent degradation of bacterial genomes is owed to a deletional bias.
The minimal genome is a concept which can be defined as the set of genes sufficient for life to exist and propagate under nutrient-rich and stress-free conditions. Alternatively, it can also be defined as the gene set supporting life on an axenic cell culture in rich media, and it is thought what makes up the minimal genome will depend on the environmental conditions that the organism inhabits. By one early investigation, the minimal genome of a bacterium should include a virtually complete set of proteins for replication and translation, a transcription apparatus including four subunits of RNA polymerase including the sigma factor rudimentary proteins sufficient for recombination and repair, several chaperone proteins, the capacity for anaerobic metabolism through glycolysis and substrate-level phosphorylation, transamination of glutamyl-tRNA to glutaminyl-tRNA, lipid biosynthesis, eight cofactor enzymes, protein export machinery, and a limited metabolite transport network including membrane ATPases. Proteins involved in the minimum bacterial genome tend to be substantially more related to proteins found in archaea and eukaryotes compared to the average gene in the bacterial genome more generally indicating a substantial number of universally conserved proteins. The minimal genomes reconstructed on the basis of existing genes does not preclude simpler systems in more primitive cells, such as an RNA world genome which does not have the need for DNA replication machinery, which is otherwise part of the minimal genome of current cells.
Nasuia deltocephalinicola was reported in 2013 to have the smallest genome of all bacteria, with 112,091 nucleotides. For comparison, the human genome has 3.2 billion nucleotides. The second smallest genome, from bacteria Tremblaya princeps, has 139,000 nucleotides. While N. deltocephalinicola has the smallest number of nucleotides, it has more protein-coding genes (137) than some bacteria.
Lokiarchaeota is a proposed phylum of the Archaea. The phylum includes all members of the group previously named Deep Sea Archaeal Group, also known as Marine Benthic Group B. Lokiarchaeota is part of the superphylum Asgard containing the phyla: Lokiarchaeota, Thorarchaeota, Odinarchaeota, Heimdallarchaeota, and Helarchaeota. A phylogenetic analysis disclosed a monophyletic grouping of the Lokiarchaeota with the eukaryotes. The analysis revealed several genes with cell membrane-related functions. The presence of such genes support the hypothesis of an archaeal host for the emergence of the eukaryotes; the eocyte-like scenarios.
A plastid is a membrane-bound organelle found in plants, algae and other eukaryotic organisms that contribute to the production of pigment molecules. Most plastids are photosynthetic, thus leading to color production and energy storage or production. There are many types of plastids in plants alone, but all plastids can be separated based on the number of times they have undergone endosymbiotic events. Currently there are three types of plastids; primary, secondary and tertiary. Endosymbiosis is reputed to have led to the evolution of eukaryotic organisms today, although the timeline is highly debated.
Novymonas esmeraldas is a protist and member of flagellated trypanosomatids. It is an obligate parasite in the gastrointestinal tract of a bug, and is in turn a host to symbiotic bacteria. It maintains strict mutualistic relationship with the bacteria as a sort of cell organelle (endosymbiont) so that it cannot lead an independent life without the bacteria. Its discovery in 2016 suggests that it is a good model in the evolution of prokaryotes into eukaryotes by symbiogenesis. The endosymbiotic bacterium was identified as member of the genus Pandoraea.
Siv Gun Elisabeth Andersson is a Swedish evolutionary biologist, professor of molecular evolution at Uppsala University. She is member of both the Royal Swedish Academy of Sciences and of Engineering. She is also Head of basic research at the Knut and Alice Wallenberg Foundation and has been co-director of the Swedish national center for large-scale research Science for Life Laboratory between 2017 and 2021. Her research focuses on the evolution of bacteria, mainly on intracellular parasites.