Reductive evolution

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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]

Contents

Examples

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]

Endosymbiotic theory

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.

Genome Sizes of Various Organisms, displaying the reduction of genomes over time to remove inessential genes. Genome Sizes.png
Genome Sizes of Various Organisms, displaying the reduction of genomes over time to remove inessential genes.

History

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.

Identification

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.

Related Research Articles

<span class="mw-page-title-main">Endosymbiont</span> Organism that lives within the body or cells of another organism

An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. 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.

<span class="mw-page-title-main">Symbiosis</span> Close, long-term biological interaction between distinct organisms (usually species)

Symbiosis is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can be either in a mutualistic, a commensalistic, or a parasitic relationship. In 1879, Heinrich Anton de Bary defined symbiosis as "the living together of unlike organisms".

<span class="mw-page-title-main">Symbiogenesis</span> Evolutionary theory holding that eukaryotic organelles evolved through symbiosis with prokaryotes

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.

<i>Rickettsia</i> Genus of bacteria

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.

<span class="mw-page-title-main">Nucleomorph</span>

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 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, obligate intracellular parasitic, aerobic bacilliform bacteria of class Alphaproteobacteria 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 semi-free-living relative of mitochondria, based on genome sequencing.

<span class="mw-page-title-main">Rickettsiales</span> Order of bacteria

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.

<span class="mw-page-title-main">Genome size</span> Amount of DNA contained in a genome

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.

<span class="mw-page-title-main">Nuclear gene</span> Gene located in the cell nucleus of a eukaryote

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.

<span class="mw-page-title-main">Cellular compartment</span> Closed part in cytosol

Cellular compartments in cell biology comprise all of the closed parts within the cytosol of a eukaryotic cell, usually surrounded by a single or double lipid layer membrane. These compartments are often, but not always, defined as membrane-bound organelles. The formation of cellular compartments is called compartmentalization.

"Candidatus Carsonella ruddii" is an obligate endosymbiotic Gammaproteobacterium with one of the smallest genomes of any characterised bacteria.

<i>Ehrlichia</i> Genus of 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.

<span class="mw-page-title-main">Prokaryote</span> Unicellular organism lacking a membrane-bound nucleus

A prokaryote is a single-celled organism whose cell lacks a nucleus and other membrane-bound organelles. The word prokaryote comes from the Ancient Greek πρό (pró), meaning 'before', and κάρυον (káruon), meaning 'nut' or 'kernel'. In the earlier two-empire system arising from the work of Édouard Chatton, prokaryotes were classified within the empire Prokaryota. However, in the three-domain system, based upon molecular phylogenetics, prokaryotes are divided into two domains: Bacteria and Archaea. A third domain, Eukaryota, consists of organisms with nuclei.

The hologenome theory of evolution recasts the individual animal or plant as a community or a "holobiont" – the host plus all of its symbiotic microbes. Consequently, the collective genomes of the holobiont form a "hologenome". Holobionts and hologenomes are structural entities that replace misnomers in the context of host-microbiota symbioses such as superorganism, organ, and metagenome. Variation in the hologenome may encode phenotypic plasticity of the holobiont and can be subject to evolutionary changes caused by selection and drift, if portions of the hologenome are transmitted between generations with reasonable fidelity. One of the important outcomes of recasting the individual as a holobiont subject to evolutionary forces is that genetic variation in the hologenome can be brought about by changes in the host genome and also by changes in the microbiome, including new acquisitions of microbes, horizontal gene transfers, and changes in microbial abundance within hosts. Although there is a rich literature on binary host–microbe symbioses, the hologenome concept distinguishes itself by including the vast symbiotic complexity inherent in many multicellular hosts.

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.

<span class="mw-page-title-main">Minimal genome</span> Concept in genetics

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 may 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.

Nasuia deltocephalinicola was reported in 2013 to have the smallest genome of all bacteria, with 112,091 nucleotides. For comparison, the genome of Escherichia coli has 4.6 million 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.

<span class="mw-page-title-main">Lokiarchaeota</span> Phylum of archaea

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.

<span class="mw-page-title-main">Plastid evolution</span> Evolution

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.

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.

References

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