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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 about 10–15% of insects.
Symbiogenesis, endosymbiotic theory, or serial endosymbiotic theory, 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 seem to be related to nitrogen-fixing filamentous cyanobacteria.
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.
Buchnera aphidicola, a member of the Pseudomonadota and the only species in the genus Buchnera, is the primary endosymbiont of aphids, and has been studied in the pea aphid, Acyrthosiphon pisum. Buchnera is believed to have had a free-living, Gram-negative ancestor similar to a modern Enterobacterales, such as Escherichia coli. Buchnera is 3 µm in diameter and has some of the key characteristics of their Enterobacterales relatives, such as a Gram-negative cell wall. However, unlike most other Gram-negative bacteria, Buchnera lacks the genes to produce lipopolysaccharides for its outer membrane. The long association with aphids and the limitation of crossover events due to strictly vertical transmission has seen the deletion of genes required for anaerobic respiration, the synthesis of amino sugars, fatty acids, phospholipids, and complex carbohydrates. This has resulted not only in one of the smallest known genomes of any living organism, but also one of the most genetically stable.
Alphaproteobacteria is a class of bacteria in the phylum Pseudomonadota. The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse and possess few commonalities, but nevertheless share a common ancestor. Like all Proteobacteria, its members are gram-negative and some of its intracellular parasitic members lack peptidoglycan and are consequently gram variable.
Bacteria are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among the first life forms to appear on Earth, and are present in most of its habitats. Bacteria inhabit soil, water, acidic hot springs, radioactive waste, and the deep biosphere of Earth's crust. Bacteria are vital in many stages of the nutrient cycle by recycling nutrients such as the fixation of nitrogen from the atmosphere. The nutrient cycle includes the decomposition of dead bodies; bacteria are responsible for the putrefaction stage in this process. In the biological communities surrounding hydrothermal vents and cold seeps, extremophile bacteria provide the nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and methane, to energy. Bacteria also live in symbiotic and parasitic relationships with plants and animals. Most bacteria have not been characterised and there are many species that cannot be grown in the laboratory. The study of bacteria is known as bacteriology, a branch of microbiology.
Acyrthosiphon pisum, commonly known as the pea aphid, is a sap-sucking insect in the family Aphididae. It feeds on several species of legumes worldwide, including forage crops, such as pea, clover, alfalfa, and broad bean, and ranks among the aphid species of major agronomical importance. The pea aphid is a model organism for biological study whose genome has been sequenced and annotated.
Liberibacter is a genus of Gram-negative bacteria in the Rhizobiaceae family. Detection of the liberibacteria is based on PCR amplification of their 16S rRNA gene with specific primers. Members of the genus are plant pathogens mostly transmitted by psyllids. The genus was originally spelled Liberobacter.
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.
Serratia symbiotica is a species of bacteria that lives as a symbiont of aphids. In the aphid Cinara cedri, it coexists with Buchnera aphidicola, given the latter cannot produce tryptophan. It is also known to habitate in Aphis fabae. Together with other endosymbionts, it provides aphids protection against parasitoids.
Hamiltonella defensa is a species of bacteria. It is maternally or sexually transmitted and lives as an endosymbiont of whiteflies and aphids, meaning that it lives within a host, protecting its host from attack. It does this through bypassing the host's immune responses by protecting its host against parasitoid wasps. However, H. defensa is only defensive if infected by a virus. H. defensa shows a relationship with Photorhabdus species, together with Regiella insecticola. Together with other endosymbionts, it provides aphids protection against parasitoids. It is known to habitate Bemisia tabaci.
Angomonas deanei is a flagellated trypanosomatid protozoan. As an obligate parasite, it infects the gastrointestinal tract of insects, and is in turn a host to symbiotic bacteria. The bacterial endosymbiont maintains a permanent mutualistic relationship with the protozoan such that it is no longer able to reproduce and survive on its own. The symbiosis, subsequently also discovered in varying degrees in other protists such as Strigomonas culicis, Novymonas esmeraldas, Diplonema japonicumand Diplonema aggregatum are considered as good models for the understanding of the evolution of eukaryotes from prokaryotes, and on the origin of cell organelles.
Sodalis is a genus of bacteria within the family Pectobacteriaceae. This genus contains several insect endosymbionts and also free-living group. It is studied due to its potential use in biological control of tsetse fly. Sodalis is important model for evolutionary biologists because of its nascent endosymbiosis with insect.
Blochmannia is a genus of symbiotic bacteria found in carpenter ant. There are over 1000 species of these ants and, as of 2014, of the over 30 species of carpenter ant that have been investigated, all contain some form of Blochmannia. The bacteria filled cells currently known as members of the genus Blochmannia were first discovered by zoologist F. Blochmann in the ovaries and midguts of insects in the 1880s. In 2000 Candidatus Blochmannia was proposed as its own genus.
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.
"Candidatus Karelsulcia muelleri" is an aerobic, gram-negative, bacillus bacterium that is a part of the phylum Bacteroidota. "Ca. K. muelleri" is an obligate and mutualistic symbiotic microbe commonly found occupying specialized cell compartments of sap-feeding insects called bacteriocytes. A majority of the research done on "Ca. K. muelleri" has detailed its relationship with the host Homalodisca vitripennis. Other studies have documented the nature of its residency in other insects like the maize leafhopper (Cicadulina) or the spittlebug (Cercopoidea). "Ca. K. muelleri" is noted for its exceptionally minimal genome and it is currently identified as having the smallest known sequenced Bacteroidota genome at only 245 kilobases.
Vertical transmission of symbionts is the transfer of a microbial symbiont from the parent directly to the offspring. Many metazoan species carry symbiotic bacteria which play a mutualistic, commensal, or parasitic role. A symbiont is acquired by a host via horizontal, vertical, or mixed transmission.
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 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. 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.
Candidatus Arsenophonus arthropodicus is a Gram-negative and intracellular secondary (S) endosymbiont that belongs to the genus Arsenophonus. This bacterium is found in the Hippoboscid louse fly, Pseudolynchia canariensis. S-endosymbionts are commonly found in distinct tissues. Strains of recovered Arsenophonus found in arthropods share 99% sequence identification in the 16S rRNA gene across all species. Arsenophonus-host interactions involve parasitism and mutualism, including a popular mechanism of "male-killing" found commonly in a related species, Arsenophonus nasoniae. This species is considered "Ca. A. arthropodicus" due it being as of yet uncultured.
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.
References
- ↑ Spaulding, A. W.; von Dohlen, C. D. (1998). "Phylogenetic Characterization and Molecular Evolution of Bacterial Endosymbionts in Psyllids (Hemiptera: Sternorrhyncha)". Molecular Biology and Evolution. 15 (11): 1506–1513. doi: 10.1093/oxfordjournals.molbev.a025878 . PMID 12572614.
- 1 2 Nakabachi A, Yamashita A, Toh H, Ishikawa H, Dunbar H, Moran N, Hattori M (2006). "The 160-kilobase genome of the bacterial endosymbiont Carsonella". Science. 314 (5797): 267. doi:10.1126/science.1134196. PMID 17038615. S2CID 44570539.
- ↑ "Taxonomy browser ("Candidatus Carsonella ruddii")". www.ncbi.nlm.nih.gov.
- ↑ Schoch, CL; Ciufo, S; Domrachev, M; Hotton, CL; Kannan, S; Khovanskaya, R; Leipe, D; Mcveigh, R; O'Neill, K; Robbertse, B; Sharma, S; Soussov, V; Sullivan, JP; Sun, L; Turner, S; Karsch-Mizrachi, I (1 January 2020). "NCBI Taxonomy: a comprehensive update on curation, resources and tools". Database: The Journal of Biological Databases and Curation. 2020. doi:10.1093/database/baaa062. PMC 7408187 . PMID 32761142.
- 1 2 Thao, MyLo L.; Moran, Nancy A.; Abbot, Patrick; Brennan, Eric B.; Burckhardt, Daniel H.; Baumann, Paul (July 2000). "Cospeciation of Psyllids and Their Primary Prokaryotic Endosymbionts". Applied and Environmental Microbiology. 66 (7): 2898–2905. Bibcode:2000ApEnM..66.2898T. doi:10.1128/aem.66.7.2898-2905.2000. PMC 92089 . PMID 10877784.
- ↑ Thao, M.L. (2001). "Phylogenetic analysis of vertically transmitted psyllid endosymbionts (Candidatus Carsonella ruddii) based on atpAGD and rpoC: comparisons with 16S-23S rDNA-derived phylogeny". Current Microbiology. 42 (6): 419–21. doi:10.1007/s002840010240. PMID 11381334. S2CID 20069209.
- ↑ Tortora(1), Funke(2), Case(3), Gerard J(1), Berdell R(2), Christine L(3) (2016). Microbiology: An Introduction (12th ed.). USA: Pearson. p. 315. ISBN 978-0321929150.
{{cite book}}: CS1 maint: multiple names: authors list (link) - ↑ Moran, Nancy A.; Bennett, Gordon M. (8 September 2014). "The Tiniest Tiny Genomes". Annual Review of Microbiology. 68 (1): 195–215. doi:10.1146/annurev-micro-091213-112901. PMID 24995872.
- ↑ Bennett, G. M.; Moran, N. A. (5 August 2013). "Small, Smaller, Smallest: The Origins and Evolution of Ancient Dual Symbioses in a Phloem-Feeding Insect". Genome Biology and Evolution. 5 (9): 1675–1688. doi:10.1093/gbe/evt118. PMC 3787670 . PMID 23918810.
- ↑ Sloan, D. B.; Moran, N. A. (19 July 2012). "Genome Reduction and Co-evolution between the Primary and Secondary Bacterial Symbionts of Psyllids". Molecular Biology and Evolution. 29 (12): 3781–3792. doi:10.1093/molbev/mss180. PMC 3494270 . PMID 22821013.