From Wikipedia the free encyclopedia

A representation of the endosymbiotic theory

An endosymbiont or endobiont[1] 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 (called rhizobia), which live in the root nodules of legumes, single-cell algae inside reef-building corals and bacterial endosymbionts that provide essential nutrients to insects.[2][3]

Endosymbiosis played key roles in the development of eukaryotes and plants. Roughly 2.2 billion years ago an archaea absorbed a bacterium through phagocytosis that eventually became the mitochondria that provide energy to all living cells. Approximately 1 billion years ago, other cells absorbed cyanobacteria that eventually became chloroplasts, organelles that produce energy from sunlight.[4] Some 100 million years ago, UCYN-A, a nitrogen-fixing bacteria became an endosymbiont of marine algae Braarudosphaera bigelowii, eventually evolving into a nitroplast.[5] Similarly, Diatoms in the family Rhopalodiaceae have evolved a diazoplast, a nitrogen-fixing organelle.[6]

Symbionts are either obligate (require their host to survive) or facultative (can survive independently).[7] The most common examples of obligate endosymbiosis are mitochondria and chloroplasts, which reproduce via mitosis in tandem with their host cells. Some human parasites, e.g. Wuchereria bancrofti and Mansonella perstans, thrive in their intermediate insect hosts because of an obligate endosymbiosis with Wolbachia spp.[8] They can both be eliminated by treatments that target their bacterial host.[9]


Endosymbiosis comes from the Greek: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living".


An overview of the endosymbiosis theory of eukaryote origin (symbiogenesis).

Symbiogenesis theory holds that eukaryotes evolved via absorbing prokaryotes. Typically, one organism envelopes a bacterium and the two evolve a mutualistic relationship. The absorbed bacteria (the endosymbiont) eventually lives exclusively within the host cells. This fits the concept of observed organelle development.[10][11][12][13][14]

Typically the endosymbiont's genome shrinks, discarding genes whose roles are displaced by the host.[15] For example the Hodgkinia genome of Magicicada cicadas is much different from the prior freestanding bacteria. The cicada life cycle involves years of stasis underground. The symbiont produces many generations during this phase, experiencing little selection pressure, allowing their genomes to diversify. Selection is episodic (when the cicadas reproduce). The original Hodgkinia genome split into three much simpler endosymbionts, each encoding only a few genes—an instance of punctuated equilibrium producing distinct lineages. The host requires all three symbionts.[16]


Symbiont transmission is the process where the host acquires its symbiont. Since symbionts are not produced by host cells, they must find their own way to reproduce and populate daughter cells as host cells divide. Horizontal, vertical, and mixed-mode (hybrid of horizonal and vertical) transmission are the three paths for symbiont transfer.


Horizontal symbiont transfer (horizontal transmission) is a process where a host acquires a facultative symbiont from the environment or another host.[7] The Rhizobia-Legume symbiosis (bacteria-plant endosymbiosis) is a prime example of this modality.[17] The Rhizobia-legume symbiotic relationship is important for processes such as the formation of root nodules. It starts with flavonoids released by the legume host, which causes the rhizobia species (endosymbiont) to activate its Nod genes.[17] These Nod genes generate lipooligosaccharide signals that the legume detects, leading to root nodule formation.[18] This process bleeds into other processes such as nitrogen fixation in plants.[17] The evolutionary advantage of such an interaction allows genetic exchange between both organisms involved to increase the propensity for novel functions as seen in the plant-bacterium interaction (holobiont formation).[19]


Vertical transmission takes place when the symbiont moves directly from parent to offspring.[20][21] In horizontal transmission each generation acquires symbionts from the environment. An example is nitrogen-fixing bacteria in certain plant roots, such as pea aphid symbionts. A third type is mixed-mode transmission, where symbionts move horizontally for some generations, after which they are acquired vertically.[22][23][24]

Wigglesworthia, a tsetse fly symbiont,[24] is vertically transmitted (via mother's milk).[25] When a symbiont reaches this stage, it resembles a cellular organelle, similar to mitochondria or chloroplasts. In vertical transmission, the symbionts do not need to survive independently, often leading them to have a reduced genome. For instance, pea aphid symbionts have lost genes for essential molecules and rely on the host to supply them. In return, the symbionts synthesize essential amino acids for the aphid host.[18] When a symbiont reaches this stage, it begins to resemble a cellular organelle, similar to mitochondria or chloroplasts. Such dependent hosts and symbionts form a holobiont. In the event of a bottleneck, a decrease in symbiont diversity could compromise host-symbiont interactions, as deleterious mutations accumulate.[26]



The best-studied examples of endosymbiosis are in invertebrates. These symbioses affect organisms with global impact, including Symbiodinium (corals), or Wolbachia (insects). Many insect agricultural pests and human disease vectors have intimate relationships with primary endosymbionts.[27]


Diagram of cospeciation, where parasites or endosymbionts speciate or branch alongside their hosts. This process is more common in hosts with primary endosymbionts.

Scientists classify insect endosymbionts as Primary or Secondary. Primary endosymbionts (P-endosymbionts) have been associated with their insect hosts for millions of years (from ten to several hundred million years). They form obligate associations and display cospeciation with their insect hosts. Secondary endosymbionts more recently associated with their hosts, may be horizontally transferred, live in the hemolymph of the insects (not specialized bacteriocytes, see below), and are not obligate.[28]


Among primary endosymbionts of insects, the best-studied are the pea aphid (Acyrthosiphon pisum) and its endosymbiont Buchnera sp. APS,[29][18] the tsetse fly Glossina morsitans morsitans and its endosymbiont Wigglesworthia glossinidia brevipalpis and the endosymbiotic protists in lower termites. As with endosymbiosis in other insects, the symbiosis is obligate. Nutritionally-enhanced diets allow symbiont-free specimens to survive, but they are unhealthy, and at best survive only a few generations.[citation needed]

In some insect groups, these endosymbionts live in specialized insect cells called bacteriocytes (also called mycetocytes), and are maternally-transmitted, i.e. the mother transmits her endosymbionts to her offspring. In some cases, the bacteria are transmitted in the egg, as in Buchnera; in others like Wigglesworthia, they are transmitted via milk to the embryo. In termites, the endosymbionts reside within the hindguts and are transmitted through trophallaxis among colony members.[30]

Primary endosymbionts are thought to help the host either by providing essential nutrients or by metabolizing insect waste products into safer forms. For example, the putative primary role of Buchnera is to synthesize essential amino acids that the aphid cannot acquire from its diet of plant sap. The primary role of Wigglesworthia is to synthesize vitamins that the tsetse fly does not get from the blood that it eats. In lower termites, the endosymbiotic protists play a major role in the digestion of lignocellulosic materials that constitute a bulk of the termites' diet.

Bacteria benefit from the reduced exposure to predators and competition from other bacterial species, the ample supply of nutrients and relative environmental stability inside the host.

Primary endosymbionts of insects have among the smallest of known bacterial genomes and have lost many genes commonly found in closely related bacteria. One theory claimed that some of these genes are not needed in the environment of the host insect cell. A complementary theory suggests that the relatively small numbers of bacteria inside each insect decrease the efficiency of natural selection in 'purging' deleterious mutations and small mutations from the population, resulting in a loss of genes over many millions of years. Research in which a parallel phylogeny of bacteria and insects was inferred supports the belief that primary endosymbionts are transferred only vertically.[31][32]

Attacking obligate bacterial endosymbionts may present a way to control their hosts, many of which are pests or human disease carriers. For example, aphids are crop pests and the tsetse fly carries the organism Trypanosoma brucei that causes African sleeping sickness.[33] Studying insect endosymbionts can aid understanding the origins of symbioses in general, as a proxy for understanding endosymbiosis in other species.

The best-studied ant endosymbionts are Blochmannia bacteria, which are the primary endosymbiont of Camponotus ants. In 2018 a new ant-associated symbiont, Candidatus Westeberhardia Cardiocondylae, was discovered in Cardiocondyla. It is reported to be a primary symbiont.[34]

Pea aphids are commonly infested by parasitic wasps. Their secondary endosymbionts attack the infesting parasitoid wasp larvae promoting the survival of both the aphid host and its endosymbionts.

The pea aphid (Acyrthosiphon pisum) contains at least three secondary endosymbionts, Hamiltonella defensa, Regiella insecticola, and Serratia symbiotica. Hamiltonella defensa defends its aphid host from parasitoid wasps.[35] This symbiosis replaces lost elements of the insect's immune response.[36]

One of the best-understood defensive symbionts is the spiral bacteria Spiroplasma poulsonii. Spiroplasma sp. can be reproductive manipulators, but also defensive symbionts of Drosophila flies. In Drosophila neotestacea, S. poulsonii has spread across North America owing to its ability to defend its fly host against nematode parasites.[37] This defence is mediated by toxins called "ribosome-inactivating proteins" that attack the molecular machinery of invading parasites.[38][39] These toxins represent one of the first understood examples of a defensive symbiosis with a mechanistic understanding for defensive symbiosis between an insect endosymbiont and its host.[40]

Sodalis glossinidius is a secondary endosymbiont of tsetse flies that lives inter- and intracellularly in various host tissues, including the midgut and hemolymph. Phylogenetic studies do not report a correlation between evolution of Sodalis and tsetse.[41] Unlike Wigglesworthia, Sodalis has been cultured in vitro.[42]

Cardinium and many other insects have secondary endosymbionts.[43][15]


Extracellular endosymbionts are represented in all four extant classes of Echinodermata (Crinoidea, Ophiuroidea, Echinoidea, and Holothuroidea). Little is known of the nature of the association (mode of infection, transmission, metabolic requirements, etc.) but phylogenetic analysis indicates that these symbionts belong to the class Alphaproteobacteria, relating them to Rhizobium and Thiobacillus. Other studies indicate that these subcuticular bacteria may be both abundant within their hosts and widely distributed among the Echinoderms.[44]

Some marine oligochaeta (e.g., Olavius algarvensis and Inanidrillus spp.) have obligate extracellular endosymbionts that fill the entire body of their host. These marine worms are nutritionally dependent on their symbiotic chemoautotrophic bacteria lacking any digestive or excretory system (no gut, mouth, or nephridia).[45]

The sea slug Elysia chlorotica's endosymbiont is the algae Vaucheria litorea. The jellyfish Mastigias have a similar relationship with an algae. Elysia chlorotica forms this relationship intracellularly with the algae's chloroplasts. These chloroplasts retain their photosynthetic capabilities and structures for several months after entering the slug's cells.[46]

Trichoplax have two bacterial endosymbionts. Ruthmannia lives inside the animal's digestive cells. Grellia lives permanently inside the endoplasmic reticulum (ER), the first known symbiont to do so.[47]

Paracatenula is a flatworm which have lived in symbiosis with an endosymbiotic bacteria for 500 million years. The bacteria produce numerous small, droplet-like vesicles that provide the host with needed nutrients.[48]


Dinoflagellate endosymbionts of the genus Symbiodinium, commonly known as zooxanthellae, are found in corals, mollusks (esp. giant clams, the Tridacna), sponges, and the unicellular foraminifera. These endosymbionts capture sunlight and provide their hosts with energy via carbonate deposition.[49]

Previously thought to be a single species, molecular phylogenetic evidence reported diversity in Symbiodinium. In some cases, the host requires a specific Symbiodinium clade. More often, however, the distribution is ecological, with symbionts switching among hosts with ease. When reefs become environmentally stressed, this distribution is related to the observed pattern of coral bleaching and recovery. Thus, the distribution of Symbiodinium on coral reefs and its role in coral bleaching is an important in coral reef ecology.[49]


In marine environments,[50][51][52][53] endosymbiont relationships are especially prevalent in oligotrophic or nutrient-poor regions of the ocean like that of the North Atlantic.[50][54][51][52] In such waters, cell growth of larger phytoplankton such as diatoms is limited by (insufficient) nitrate concentrations.[55]  Endosymbiotic bacteria fix nitrogen for their hosts and in turn receive organic carbon from photosynthesis.[54] These symbioses play an important role in global carbon cycling.[56][51][52]

One known symbiosis between the diatom Hemialus spp. and the cyanobacterium Richelia intracellularis has been reported in North Atlantic, Mediterranean, and Pacific waters.[50][51][57] Richelia is found within the diatom frustule of Hemiaulus spp., and has a reduced genome.[58] A 2011 study measured nitrogen fixation by the cyanobacterial host Richelia intracellularis well above intracellular requirements, and found the cyanobacterium was likely fixing nitrogen for its host.[55] Additionally, both host and symbiont cell growth were much greater than free-living Richelia intracellularis or symbiont-free Hemiaulus spp.[55] The Hemaiulus-Richelia symbiosis is not obligatory, especially in nitrogen-replete areas.[50]

Richelia intracellularis is also found in Rhizosolenia spp., a diatom found in oligotrophic oceans.[54][55][52] Compared to the Hemaiulus host, the endosymbiosis with Rhizosolenia is much more consistent, and Richelia intracellularis is generally found in Rhizosolenia.[50] There are some asymbiotic (occurs without an endosymbiont) Rhizosolenia, however there appears to be mechanisms limiting growth of these organisms in low nutrient conditions.[59] Cell division for both the diatom host and cyanobacterial symbiont can be uncoupled and mechanisms for passing bacterial symbionts to daughter cells during cell division are still relatively unknown.[59]

Other endosymbiosis with nitrogen fixers in open oceans include Calothrix in Chaetoceros spp. and UNCY-A in prymnesiophyte microalga.[60]  The Chaetoceros-Calothrix endosymbiosis is hypothesized to be more recent, as the Calothrix genome is generally intact. While other species like that of the UNCY-A symbiont and Richelia have reduced genomes.[58] This reduction in genome size occurs within nitrogen metabolism pathways indicating endosymbiont species are generating nitrogen for their hosts and losing the ability to use this nitrogen independently.[58] This endosymbiont reduction in genome size, might be a step that occurred in the evolution of organelles (above).[60]


Mixotricha paradoxa is a protozoan that lacks mitochondria. However, spherical bacteria live inside the cell and serve the function of the mitochondria. Mixotricha has three other species of symbionts that live on the surface of the cell.[61]

Paramecium bursaria, a species of ciliate, has a mutualistic symbiotic relationship with green alga called Zoochlorella. The algae live in its cytoplasm.[62]

Platyophrya chlorelligera is a freshwater ciliate that harbors Chlorella that perform photosynthesis.[63][64]

Strombidium purpureum is a marine ciliate that uses endosymbiotic, purple, non-sulphur bacteria for anoxygenic photosynthesis.[65][66]

Paulinella chromatophora is a freshwater amoeboid that has a cyanobacterium endosymbiont.

Many foraminifera are hosts to several types of algae, such as red algae, diatoms, dinoflagellates and chlorophyta.[67] These endosymbionts can be transmitted vertically to the next generation via asexual reproduction of the host, but because the endosymbionts are larger than the foraminiferal gametes, they need to acquire algae horizontally following sexual reproduction.[68]

Several species of radiolaria have photosynthetic symbionts. In some species the host digests algae to keep the population at a constant level.[69]

Hatena arenicola is a flagellate protist with a complicated feeding apparatus that feeds on other microbes. When it engulfs a green Nephroselmis alga, the feeding apparatus disappears and it becomes photosynthetic. During mitosis the algae is transferred to only one of the daughter cells, while the other cell restarts the cycle.

In 1966, biologist Kwang W. Jeon found that a lab strain of Amoeba proteus had been infected by bacteria that lived inside the cytoplasmic vacuoles.[70] This infection killed almost all of the infected protists. After the equivalent of 40 host generations, the two organisms become mutually interdependent. A genetic exchange between the prokaryotes and protists occurred.[71][72][73]


The spotted salamander (Ambystoma maculatum) lives in a relationship with the algae Oophila amblystomatis, which grows in its egg cases.[74]


All vascular plants harbor endosymbionts or endophytes in this context. They include bacteria, fungi, viruses, protozoa and even microalgae. Endophytes aid in processes such as growth and development, nutrient uptake, and defense against biotic and abiotic stresses like drought, salinity, heat, and herbivores.[75]

Plant symbionts can be categorized into epiphytic, endophytic, and mycorrhizal. These relations can also be categorized as beneficial, mutualistic, neutral, and pathogenic.[76][77] Microorganisms living as endosymbionts in plants can enhance their host's primary productivity either by producing or capturing important resources.[78] These endosymbionts can also enhance plant productivity by producing toxic metabolites that aid plant defenses against herbivores.[79][80]

Plants are dependent on plastid or chloroplast organelles. The chloroplast is derived from a cyanobacterial primary endosymbiosis that began over one billion years ago. An oxygenic, photosynthetic free-living cyanobacterium was engulfed and kept by a heterotrophic protist and eventually evolved into the present intracellular organelle.[81]  

Mycorrhizal endosymbionts appear only in fungi.

Typically, plant endosymbiosis studies focus on a single category or species to better understand their individual biological processes and functions.[82]

Fungal endophytes[edit]

Fungal endophytes can be found in all plant tissues. Fungi living below the ground amidst plant roots are known as mycorrhiza, but are further categorized based on their location inside the root, with prefixes such as ecto, endo, arbuscular, ericoid, etc. Fungal endosymbionts that live in the roots and extend their extraradical hyphae into the outer rhizosphere are known as ectendosymbionts.[83][84]

Arbuscular Mycorrhizal Fungi (AMF)[edit]

Arbuscular mycorrhizal fungi or AMF are the most diverse plant microbial endosymbionts. With exceptions such as the Ericaceae family, almost all vascular plants harbor AMF endosymbionts as endo and ecto as well. AMF plant endosymbionts systematically colonize plant roots and help the plant host acquire soil nutrients such as nitrogen. In return it absorbs plant organic carbon products.[83] Plant root exudates contain diverse secondary metabolites, especially flavonoids and strigolactones that act as chemical signals and attracts the AMF.[85] AMF Gigaspora margarita lives as a plant endosymbiont and also harbors further endosymbiont intracytoplasmic bacterium-like organisms.[86] AMF generally promote plant health and growth and alleviate abiotic stresses such as salinity, drought, heat, poor nutrition, and metal toxicity.[87] Individual AMF species have different effects in different hosts – introducing the AMF of one plant to another plant can reduce the latter's growth.[88]

Endophytic fungi[edit]

Endophytic fungi in mutualistic relations directly benefit and benefit from their host plants. They also can help their hosts succeed in polluted environments such as those contaminated with toxic metals.[89] Fungal endophytes are taxonomically diverse and are divided into categories based on mode of transmission, biodiversity, in planta colonization and host plant type.[90][91] Clavicipitaceous fungi systematically colonize temperate season grasses. Non-clavicipitaceous fungi colonize higher plants and even roots and divide into subcategories.[92] Aureobasidium and preussia species of endophytic fungi isolated from Boswellia sacra produce indole acetic acid hormone to promote plant health and development.[93]

Aphids can be found in most plants. Carnivorous ladybirds are aphid predators and are used in pest control. Plant endophytic fungus Neotyphodium lolii produces alkaloid mycotoxins in response to aphid invasions. In response, ladybird predators exhibited reduced fertility and abnormal reproduction, suggesting that the mycotoxins are transmitted along the food chain and affect the predators.[78]

Endophytic bacteria[edit]

Endophytic bacteria belong to a diverse group of plant endosymbionts characterized by systematic colonization of plant tissues. The most common genera include Pseudomonas, Bacillus, Acinetobacter, Actinobacteria, Sphingomonas. Some endophytic bacteria, such as Bacillus amyloliquefaciens, a seed-born endophytic bacteria, produce plant growth by producing gibberellins, which are potent plant growth hormones. Bacillus amyloliquefaciens promotes the taller height of transgenic dwarf rice plants.[94] Some endophytic bacteria genera additionally belong to the Enterobacteriaceae family.[95] Endophytic bacteria typically colonize the leaf tissues from plant roots, but can also enter the plant through the leaves through leaf stomata.[96] Generally, the endophytic bacteria are isolated from the plant tissues by surface sterilization of the plant tissue in a sterile environment.[97] Passenger endophytic bacteria eventually colonize inner tissue of plant by stochastic events while True endophytes possess adaptive traits because of which they live strictly in association with plants.[98] The in vitro-cultivated endophytic bacteria association with plants is considered a more intimate relationship that helps plants acclimatize to conditions and promotes health and growth. Endophytic bacteria are considered to be plant's essential endosymbionts because virtually all plants harbor them, and these endosymbionts play essential roles in host survival.[99] This endosymbiotic relation is important in terms of ecology, evolution and diversity. Endophytic bacteria such as Sphingomonas sp. and Serratia sp. that are isolated from arid land plants regulate endogenous hormone content and promote growth.[100]

Archaea endosymbionts[edit]

Archaea are members of most microbiomes. While archaea are abundant in extreme environments, they are less abundant and diverse in association with eukaryotic hosts. Nevertheless, archaea are a substantial constituent of plant-associated ecosystems in the aboveground and belowground phytobiome, and play a role in host plant’s health, growth and survival amid biotic and abiotic stresses. However, few studies have investigated the role of archaea in plant health and its symbiotic relationships.[101] Most plant endosymbiosis studies focus on fungal or bacteria using metagenomic approaches.[102]

The characterization of archaea includes crop plants such as rice[103] and maize, but also aquatic plants.[101] The abundance of archaea varies by tissue type; for example archaea are more abundant in the rhizosphere than the phyllosphere and endosphere.[104] This archaeal abundance is associated with plant species type, environment and the plant’s developmental stage.[105] In a study on plant genotype-specific archaeal and bacterial endophytes, 35% of archaeal sequences were detected in overall sequences (achieved using amplicon sequencing and verified by real time-PCR). The archaeal sequences belong to the phyla Thaumarchaeota, Crenarchaeota, and Euryarchaeota.[106]


Some Betaproteobacteria have Gammaproteobacteria endosymbionts.[107]


Fungi host endohyphal bacteria;[108] the effects of the bacteria are not well studied. Many such fungi in turn live within plants.[108] These fungi are otherwise known as fungal endophytes. It is hypothesized that the fungi offers a safe haven for the bacteria, and the diverse bacteria that they attract create a micro-ecosystem.[109]

These interactions may impact the way that fungi interact with the environment by modulating their phenotypes.[108] The bacteria do this by altering the fungi's gene expression.[108] For example, Luteibacter sp. has been shown to naturally infect the ascomycetous endophyte Pestalotiopsis sp. isolated from Platycladus orientalis.[108] The Luteibacter sp. influences the auxin and enzyme production within its host, which, in turn, may influence the effect the fungus has on its plant host.[108] Another interesting example of a bacteria living in symbiosis with a fungus is the fungus Mortierella. This soil-dwelling fungus lives in close association with a toxin-producing bacteria, Mycoavidus, which helps the fungus defend against nematodes.[110]

Virus endosymbionts[edit]

The human genome project found several thousand endogenous retroviruses, endogenous viral elements in the genome that closely resemble and can be derived from retroviruses, organized into 24 families.[111][citation needed][112]

See also[edit]


  1. ^ Margulis L, Chapman MJ (2009). Kingdoms & domains an illustrated guide to the phyla of life on Earth (4th ed.). Amsterdam: Academic Press/Elsevier. p. 493. ISBN 978-0-08-092014-6.
  2. ^ Mergaert P (April 2018). "Role of antimicrobial peptides in controlling symbiotic bacterial populations". Natural Product Reports. 35 (4): 336–356. doi:10.1039/c7np00056a. PMID 29393944.
  3. ^ Little AF, van Oppen MJ, Willis BL (June 2004). "Flexibility in algal endosymbioses shapes growth in reef corals". Science. 304 (5676): 1492–1494. Bibcode:2004Sci...304.1491L. doi:10.1126/science.1095733. PMID 15178799. S2CID 10050417.
  4. ^ Baisas, Laura (18 April 2024). "For the first time in one billion years, two lifeforms truly merged into one organism". Popular Science. Retrieved 26 April 2024.
  5. ^ Wong, Carissa (11 April 2024). "Scientists discover first algae that can fix nitrogen — thanks to a tiny cell structure". Nature.com. Archived from the original on 14 April 2024. Retrieved 16 April 2024.
  6. ^ "Genomic divergence within non-photosynthetic cyanobacterial endosymbionts in rhopalodiacean diatoms". www.ncbi.nlm.nih.gov. PMC 5638926. PMID 29026213. Retrieved 25 April 2024.
  7. ^ a b Bright, Monika; Bulgheresi, Silvia (March 2010). "A complex journey: transmission of microbial symbionts". Nature Reviews Microbiology. 8 (3): 218–230. doi:10.1038/nrmicro2262. ISSN 1740-1534. PMC 2967712. PMID 20157340.
  8. ^ Slatko, Barton E.; Taylor, Mark J.; Foster, Jeremy M. (1 July 2010). "The Wolbachia endosymbiont as an anti-filarial nematode target". Symbiosis. 51 (1): 55–65. Bibcode:2010Symbi..51...55S. doi:10.1007/s13199-010-0067-1. ISSN 1878-7665. PMC 2918796. PMID 20730111.
  9. ^ Warrell D, Cox TM, Firth J, Török E (11 October 2012). Oxford Textbook of Medicine: Infection. OUP Oxford. ISBN 978-0-19-965213-6.
  10. ^ Moore KR, Magnabosco C, Momper L, Gold DA, Bosak T, Fournier GP (2019). "An Expanded Ribosomal Phylogeny of Cyanobacteria Supports a Deep Placement of Plastids". Frontiers in Microbiology. 10: 1612. doi:10.3389/fmicb.2019.01612. PMC 6640209. PMID 31354692.
  11. ^ McCutcheon JP (October 2021). "The Genomics and Cell Biology of Host-Beneficial Intracellular Infections". Annual Review of Cell and Developmental Biology. 37 (1): 115–142. doi:10.1146/annurev-cellbio-120219-024122. PMID 34242059. S2CID 235786110.
  12. ^ Callier V (8 June 2022). "Mitochondria and the origin of eukaryotes". Knowable Magazine. doi:10.1146/knowable-060822-2. Retrieved 18 August 2022.
  13. ^ Sagan L (March 1967). "On the origin of mitosing cells". Journal of Theoretical Biology. 14 (3): 255–274. Bibcode:1967JThBi..14..225S. doi:10.1016/0022-5193(67)90079-3. PMID 11541392.
  14. ^ Gabaldón T (October 2021). "Origin and Early Evolution of the Eukaryotic Cell". Annual Review of Microbiology. 75 (1): 631–647. doi:10.1146/annurev-micro-090817-062213. PMID 34343017. S2CID 236916203.
  15. ^ a b Wernegreen JJ (November 2002). "Genome evolution in bacterial endosymbionts of insects". Nature Reviews. Genetics. 3 (11): 850–861. doi:10.1038/nrg931. PMID 12415315. S2CID 29136336.
  16. ^ Campbell MA, Łukasik P, Simon C, McCutcheon JP (November 2017). "Idiosyncratic Genome Degradation in a Bacterial Endosymbiont of Periodical Cicadas". Current Biology. 27 (22): 3568–3575.e3. Bibcode:2017CBio...27E3568C. doi:10.1016/j.cub.2017.10.008. PMC 8879801. PMID 29129532.
  17. ^ a b c Gage, Daniel J. (June 2004). "Infection and Invasion of Roots by Symbiotic, Nitrogen-Fixing Rhizobia during Nodulation of Temperate Legumes". Microbiology and Molecular Biology Reviews. 68 (2): 280–300. doi:10.1128/MMBR.68.2.280-300.2004. ISSN 1092-2172. PMC 419923. PMID 15187185.
  18. ^ a b c Shigenobu S, Watanabe H, Hattori M, Sakaki Y, Ishikawa H (September 2000). "Genome sequence of the endocellular bacterial symbiont of aphids Buchnera sp. APS". Nature. 407 (6800): 81–86. Bibcode:2000Natur.407...81S. doi:10.1038/35024074. PMID 10993077.
  19. ^ Chrostek, Ewa; Pelz-Stelinski, Kirsten; Hurst, Gregory D. D.; Hughes, Grant L. (2017). "Horizontal Transmission of Intracellular Insect Symbionts via Plants". Frontiers in Microbiology. 8: 2237. doi:10.3389/fmicb.2017.02237. ISSN 1664-302X. PMC 5712413. PMID 29234308.
  20. ^ McCutcheon JP (October 2021). "The Genomics and Cell Biology of Host-Beneficial Intracellular Infections". Annual Review of Cell and Developmental Biology. 37 (1): 115–142. doi:10.1146/annurev-cellbio-120219-024122. PMID 34242059. S2CID 235786110.
  21. ^ Callier V (8 June 2022). "Mitochondria and the origin of eukaryotes". Knowable Magazine. doi:10.1146/knowable-060822-2. Retrieved 18 August 2022.
  22. ^ Wierz JC, Gaube P, Klebsch D, Kaltenpoth M, Flórez LV (2021). "Transmission of Bacterial Symbionts With and Without Genome Erosion Between a Beetle Host and the Plant Environment". Frontiers in Microbiology. 12: 715601. doi:10.3389/fmicb.2021.715601. PMC 8493222. PMID 34630349.
  23. ^ Ebert D (23 November 2013). "The Epidemiology and Evolution of Symbionts with Mixed-Mode Transmission". Annual Review of Ecology, Evolution, and Systematics. 44 (1): 623–643. doi:10.1146/annurev-ecolsys-032513-100555. ISSN 1543-592X. Retrieved 19 August 2022.
  24. ^ a b Bright M, Bulgheresi S (March 2010). "A complex journey: transmission of microbial symbionts". Nature Reviews. Microbiology. 8 (3): 218–230. doi:10.1038/nrmicro2262. PMC 2967712. PMID 20157340.
  25. ^ Bright M, Bulgheresi S (March 2010). "A complex journey: transmission of microbial symbionts". Nature Reviews. Microbiology. 8 (3): 218–230. doi:10.1038/nrmicro2262. PMC 2967712. PMID 20157340.
  26. ^ Smith, Noel H.; Gordon, Stephen V.; de la Rua-Domenech, Ricardo; Clifton-Hadley, Richard S.; Hewinson, R. Glyn (September 2006). "Bottlenecks and broomsticks: the molecular evolution of Mycobacterium bovis". Nature Reviews Microbiology. 4 (9): 670–681. doi:10.1038/nrmicro1472. ISSN 1740-1534. PMID 16912712. S2CID 2015074.
  27. ^ Eleftherianos, Ioannis; Atri, Jaishri; Accetta, Julia; Castillo, Julio C. (2013). "Endosymbiotic bacteria in insects: guardians of the immune system?". Frontiers in Physiology. 4: 46. doi:10.3389/fphys.2013.00046. ISSN 1664-042X. PMC 3597943. PMID 23508299.
  28. ^ Baumann P, Moran NA, Baumann L (2000). "Bacteriocyte-associated endosymbionts of insects". In Dworkin M (ed.). The prokaryotes. New York: Springer.
  29. ^ Douglas AE (January 1998). "Nutritional interactions in insect-microbial symbioses: aphids and their symbiotic bacteria Buchnera". Annual Review of Entomology. 43: 17–37. doi:10.1146/annurev.ento.43.1.17. PMID 15012383. S2CID 29594533.
  30. ^ Nalepa, Christine A. (2020). "Origin of Mutualism Between Termites and Flagellated Gut Protists: Transition From Horizontal to Vertical Transmission". Frontiers in Ecology and Evolution. 8. doi:10.3389/fevo.2020.00014. ISSN 2296-701X.
  31. ^ Wernegreen JJ (March 2004). "Endosymbiosis: lessons in conflict resolution". PLOS Biology. 2 (3): E68. doi:10.1371/journal.pbio.0020068. PMC 368163. PMID 15024418.
  32. ^ Moran NA (April 1996). "Accelerated evolution and Muller's rachet in endosymbiotic bacteria". Proceedings of the National Academy of Sciences of the United States of America. 93 (7): 2873–2878. Bibcode:1996PNAS...93.2873M. doi:10.1073/pnas.93.7.2873. PMC 39726. PMID 8610134.
  33. ^ Aksoy S, Maudlin I, Dale C, Robinson AS, O'Neill SL (January 2001). "Prospects for control of African trypanosomiasis by tsetse vector manipulation". Trends in Parasitology. 17 (1): 29–35. doi:10.1016/S1471-4922(00)01850-X. PMID 11137738.
  34. ^ Klein A, Schrader L, Gil R, Manzano-Marín A, Flórez L, Wheeler D, et al. (February 2016). "A novel intracellular mutualistic bacterium in the invasive ant Cardiocondyla obscurior". The ISME Journal. 10 (2): 376–388. Bibcode:2016ISMEJ..10..376K. doi:10.1038/ismej.2015.119. PMC 4737929. PMID 26172209.
  35. ^ Oliver KM, Campos J, Moran NA, Hunter MS (February 2008). "Population dynamics of defensive symbionts in aphids". Proceedings. Biological Sciences. 275 (1632): 293–299. doi:10.1098/rspb.2007.1192. PMC 2593717. PMID 18029301.
  36. ^ International Aphid Genomics Consortium (February 2010). "Genome sequence of the pea aphid Acyrthosiphon pisum". PLOS Biology. 8 (2): e1000313. doi:10.1371/journal.pbio.1000313. PMC 2826372. PMID 20186266.
  37. ^ Jaenike J, Unckless R, Cockburn SN, Boelio LM, Perlman SJ (July 2010). "Adaptation via symbiosis: recent spread of a Drosophila defensive symbiont". Science. 329 (5988): 212–215. Bibcode:2010Sci...329..212J. doi:10.1126/science.1188235. PMID 20616278. S2CID 206526012.
  38. ^ Hamilton PT, Peng F, Boulanger MJ, Perlman SJ (January 2016). "A ribosome-inactivating protein in a Drosophila defensive symbiont". Proceedings of the National Academy of Sciences of the United States of America. 113 (2): 350–355. Bibcode:2016PNAS..113..350H. doi:10.1073/pnas.1518648113. PMC 4720295. PMID 26712000.
  39. ^ Ballinger MJ, Perlman SJ (July 2017). "Generality of toxins in defensive symbiosis: Ribosome-inactivating proteins and defense against parasitic wasps in Drosophila". PLOS Pathogens. 13 (7): e1006431. doi:10.1371/journal.ppat.1006431. PMC 5500355. PMID 28683136.
  40. ^ Ballinger MJ, Perlman SJ (July 2017). "Generality of toxins in defensive symbiosis: Ribosome-inactivating proteins and defense against parasitic wasps in Drosophila". PLOS Pathogens. 13 (7): e1006431. doi:10.1371/journal.ppat.1006431. PMC 5500355. PMID 28683136.
  41. ^ Aksoy, S., Pourhosseini, A. & Chow, A. 1995. Mycetome endosymbionts of tsetse flies constitute a distinct lineage related to Enterobacteriaceae. Insect Mol Biol. 4, 15–22.
  42. ^ Welburn SC, Maudlin I, Ellis DS (June 1987). "In vitro cultivation of rickettsia-like-organisms from Glossina spp". Annals of Tropical Medicine and Parasitology. 81 (3): 331–335. doi:10.1080/00034983.1987.11812127. PMID 3662675.
  43. ^ Zchori-Fein E, Perlman SJ (July 2004). "Distribution of the bacterial symbiont Cardinium in arthropods". Molecular Ecology. 13 (7): 2009–2016. Bibcode:2004MolEc..13.2009Z. doi:10.1111/j.1365-294X.2004.02203.x. PMID 15189221. S2CID 24361903.
  44. ^ Burnett WJ, McKenzie JD (May 1997). "Subcuticular bacteria from the brittle star Ophiactis balli (Echinodermata: Ophiuroidea) represent a new lineage of extracellular marine symbionts in the alpha subdivision of the class Proteobacteria". Applied and Environmental Microbiology. 63 (5): 1721–1724. Bibcode:1997ApEnM..63.1721B. doi:10.1128/AEM.63.5.1721-1724.1997. PMC 168468. PMID 9143108.
  45. ^ Dubilier N, Mülders C, Ferdelman T, de Beer D, Pernthaler A, Klein M, et al. (May 2001). "Endosymbiotic sulphate-reducing and sulphide-oxidizing bacteria in an oligochaete worm". Nature. 411 (6835): 298–302. Bibcode:2001Natur.411..298D. doi:10.1038/35077067. PMID 11357130. S2CID 4420931.
  46. ^ Mujer CV, Andrews DL, Manhart JR, Pierce SK, Rumpho ME (October 1996). "Chloroplast genes are expressed during intracellular symbiotic association of Vaucheria litorea plastids with the sea slug Elysia chlorotica". Proceedings of the National Academy of Sciences of the United States of America. 93 (22): 12333–12338. Bibcode:1996PNAS...9312333M. doi:10.1073/pnas.93.22.12333. PMC 37991. PMID 8901581.
  47. ^ Society, Max Planck. "Deceptively simple: Minute marine animals live in a sophisticated symbiosis with bacteria". phys.org.
  48. ^ Society, Max Planck. "How a bacterium feeds an entire flatworm". phys.org.
  49. ^ a b Baker AC (November 2003). "Flexibility and Specificity in Coral-Algal Symbiosis: Diversity, Ecology, and Biogeography of Symbiodinium". Annual Review of Ecology, Evolution, and Systematics. 34: 661–89. doi:10.1146/annurev.ecolsys.34.011802.132417. S2CID 35278104.
  50. ^ a b c d e Villareal T (1994). "Widespread occurrence of the Hemiaulus-cyanobacterial symbiosis in the southwest North Atlantic Ocean". Bulletin of Marine Science. 54: 1–7.
  51. ^ a b c d Carpenter EJ, Montoya JP, Burns J, Mulholland MR, Subramaniam A, Capone DG (20 August 1999). "Extensive bloom of a N2-fixing diatom/cyanobacterial association in the tropical Atlantic Ocean". Marine Ecology Progress Series. 185: 273–283. Bibcode:1999MEPS..185..273C. doi:10.3354/meps185273. hdl:1853/43100.
  52. ^ a b c d Foster RA, Subramaniam A, Mahaffey C, Carpenter EJ, Capone DG, Zehr JP (March 2007). "Influence of the Amazon River plume on distributions of free-living and symbiotic cyanobacteria in the western tropical north Atlantic Ocean". Limnology and Oceanography. 52 (2): 517–532. Bibcode:2007LimOc..52..517F. doi:10.4319/lo.2007.52.2.0517. S2CID 53504106.
  53. ^ Subramaniam A, Yager PL, Carpenter EJ, Mahaffey C, Björkman K, Cooley S, et al. (July 2008). "Amazon River enhances diazotrophy and carbon sequestration in the tropical North Atlantic Ocean". Proceedings of the National Academy of Sciences of the United States of America. 105 (30): 10460–10465. doi:10.1073/pnas.0710279105. PMC 2480616. PMID 18647838.
  54. ^ a b c Goebel NL, Turk KA, Achilles KM, Paerl R, Hewson I, Morrison AE, et al. (December 2010). "Abundance and distribution of major groups of diazotrophic cyanobacteria and their potential contribution to N₂ fixation in the tropical Atlantic Ocean". Environmental Microbiology. 12 (12): 3272–3289. Bibcode:2010EnvMi..12.3272G. doi:10.1111/j.1462-2920.2010.02303.x. PMID 20678117.
  55. ^ a b c d Foster RA, Kuypers MM, Vagner T, Paerl RW, Musat N, Zehr JP (September 2011). "Nitrogen fixation and transfer in open ocean diatom-cyanobacterial symbioses". The ISME Journal. 5 (9): 1484–1493. Bibcode:2011ISMEJ...5.1484F. doi:10.1038/ismej.2011.26. PMC 3160684. PMID 21451586.
  56. ^ Scharek R, Tupas LM, Karl DM (11 June 1999). "Diatom fluxes to the deep sea in the oligotrophic North Pacific gyre at Station Aloha". Marine Ecology Progress Series. 182: 55–67. Bibcode:1999MEPS..182...55S. doi:10.3354/meps182055. hdl:10261/184131.
  57. ^ Zeev EB, Yogev T, Man-Aharonovich D, Kress N, Herut B, Béjà O, Berman-Frank I (September 2008). "Seasonal dynamics of the endosymbiotic, nitrogen-fixing cyanobacterium Richelia intracellularis in the eastern Mediterranean Sea". The ISME Journal. 2 (9): 911–923. Bibcode:2008ISMEJ...2..911Z. doi:10.1038/ismej.2008.56. PMID 18580972.
  58. ^ a b c Hilton JA, Foster RA, Tripp HJ, Carter BJ, Zehr JP, Villareal TA (23 April 2013). "Genomic deletions disrupt nitrogen metabolism pathways of a cyanobacterial diatom symbiont". Nature Communications. 4 (1): 1767. Bibcode:2013NatCo...4.1767H. doi:10.1038/ncomms2748. PMC 3667715. PMID 23612308.
  59. ^ a b Villareal TA (December 1989). "Division cycles in the nitrogen-fixingRhizosolenia(Bacillariophyceae)-Richelia(Nostocaceae) symbiosis". British Phycological Journal. 24 (4): 357–365. doi:10.1080/00071618900650371.
  60. ^ a b Zehr JP (September 2015). "EVOLUTION. How single cells work together". Science. 349 (6253): 1163–1164. doi:10.1126/science.aac9752. PMID 26359387. S2CID 206641230.
  61. ^ Wenzel, Marika; Radek, Renate; Brugerolle, Guy; König, Helmut (1 January 2003). "Identification of the ectosymbiotic bacteria of Mixotricha paradoxa involved in movement symbiosis". European Journal of Protistology. 39 (1): 11–23. doi:10.1078/0932-4739-00893. ISSN 0932-4739.
  62. ^ Dziallas, C.; Allgaier, M.; Monaghan, M. T.; Grossart, H. P. (2012). "Act together—implications of symbioses in aquatic ciliates". Frontiers in Microbiology. 3: 288. doi:10.3389/fmicb.2012.00288. PMC 3413206. PMID 22891065.
  63. ^ Joint, Ian (29 June 2013). Molecular Ecology of Aquatic Microbes. Springer Science & Business Media. ISBN 978-3-642-79923-5.
  64. ^ Kawakami, H. (1991). "An endosymbiotic Chlorella-bearing ciliate: Platyophrya chlorelligera Kawakami 1989". European Journal of Protistology. 26 (3–4): 245–255. doi:10.1016/S0932-4739(11)80146-X. PMID 23196282.
  65. ^ Fenchel, Tom; Bernard, Catherine (1993). "Endosymbiotic purple non-sulphur bacteria in an anaerobic ciliated protozoon". FEMS Microbiology Letters. 110: 21–25. doi:10.1111/j.1574-6968.1993.tb06289.x. S2CID 86458030.
  66. ^ Paracer, Surindar; Ahmadjian, Vernon (6 July 2000). Symbiosis: An Introduction to Biological Associations. Oxford University Press. ISBN 978-0-19-802788-1.
  67. ^ Joseph Seckbach; Patrick Kociolek (2011). The Diatom World. Springer Science & Business Media. p. 439. ISBN 978-94-007-1327-7.
  68. ^ Toledo, Rafael Isaac Ponce (5 March 2018). Origins and early evolution of photosynthetic eukaryotes (Thesis). Université Paris-Saclay. S2CID 89705815.
  69. ^ Surindar Paracer; Vernon Ahmadjian (2000). Symbiosis: An Introduction to Biological Associations. Oxford University Press. p. 155. ISBN 978-0-19-511807-0.
  70. ^ Jeon KW, Jeon MS (October 1976). "Endosymbiosis in amoebae: recently established endosymbionts have become required cytoplasmic components". Journal of Cellular Physiology. 89 (2): 337–344. doi:10.1002/jcp.1040890216. PMID 972171. S2CID 32044949.
  71. ^ "Kwang W. Jeon | Biochemistry & Cellular and Molecular Biology – UTK BCMB". 28 April 2014. Archived from the original on 31 August 2018. Retrieved 14 May 2019.
  72. ^ Luigi Nibali; Brian Henderson (2016). The Human Microbiota and Chronic Disease: Dysbiosis as a Cause of Human Pathology. John Wiley & Sons. p. 165. ISBN 978-1-118-98287-7.
  73. ^ K. Jeon, “Amoeba and X-bacteria: Symbiont Acquisition and Possible Species Change,” in: L. Margulis and R. Fester, eds., Symbiosis as a Source of Evolutionary Innovation (Cambridge, Mass.: MIT Press), c. 9.
  74. ^ Kerney R, Kim E, Hangarter RP, Heiss AA, Bishop CD, Hall BK (April 2011). "Intracellular invasion of green algae in a salamander host". Proceedings of the National Academy of Sciences of the United States of America. 108 (16): 6497–6502. Bibcode:2011PNAS..108.6497K. doi:10.1073/pnas.1018259108. PMC 3080989. PMID 21464324.
  75. ^ Baron NC, Rigobelo EC (2022). "Endophytic fungi: a tool for plant growth promotion and sustainable agriculture". Mycology. 13 (1): 39–55. doi:10.1080/21501203.2021.1945699. PMC 8856089. PMID 35186412.
  76. ^ Hardoim PR, van Overbeek LS, Berg G, Pirttilä AM, Compant S, Campisano A, et al. (September 2015). "The Hidden World within Plants: Ecological and Evolutionary Considerations for Defining Functioning of Microbial Endophytes". Microbiology and Molecular Biology Reviews. 79 (3): 293–320. doi:10.1128/MMBR.00050-14. PMC 4488371. PMID 26136581.
  77. ^ Khare E, Mishra J, Arora NK (2018). "Multifaceted Interactions Between Endophytes and Plant: Developments and Prospects". Frontiers in Microbiology. 9: 2732. doi:10.3389/fmicb.2018.02732. PMC 6249440. PMID 30498482.
  78. ^ a b de Sassi C, Müller CB, Krauss J (May 2006). "Fungal plant endosymbionts alter life history and reproductive success of aphid predators". Proceedings. Biological Sciences. 273 (1591): 1301–1306. doi:10.1098/rspb.2005.3442. PMC 1560287. PMID 16720406.
  79. ^ Schardl CL, Leuchtmann A, Spiering MJ (2 June 2004). "Symbioses of grasses with seedborne fungal endophytes". Annual Review of Plant Biology. 55 (1): 315–340. doi:10.1146/annurev.arplant.55.031903.141735. PMID 15377223.
  80. ^ Hunter MD, Price PW (1992). "Playing Chutes and Ladders: Heterogeneity and the Relative Roles of Bottom-Up and Top-Down Forces in Natural Communities". Ecology. 73 (3): 724–732. Bibcode:1992Ecol...73..724H. doi:10.2307/1940152. ISSN 0012-9658. JSTOR 1940152. S2CID 54005488.
  81. ^ Qiu H, Yoon HS, Bhattacharya D (September 2013). "Algal endosymbionts as vectors of horizontal gene transfer in photosynthetic eukaryotes". Frontiers in Plant Science. 4: 366. doi:10.3389/fpls.2013.00366. PMC 3777023. PMID 24065973.
  82. ^ Porras-Alfaro A, Bayman P (8 September 2011). "Hidden fungi, emergent properties: endophytes and microbiomes". Annual Review of Phytopathology. 49 (1): 291–315. doi:10.1146/annurev-phyto-080508-081831. PMID 19400639.
  83. ^ a b Salhi LN, Bustamante Villalobos P, Forget L, Burger G, Lang BF (September 2022). "Endosymbionts in cranberry: Diversity, effect on plant growth, and pathogen biocontrol". Plants, People, Planet. 4 (5): 511–522. doi:10.1002/ppp3.10290. ISSN 2572-2611. S2CID 250548548.
  84. ^ Roth R, Paszkowski U (October 2017). "Plant carbon nourishment of arbuscular mycorrhizal fungi". Current Opinion in Plant Biology. 39 Cell signalling and gene regulation 2017. 39: 50–56. Bibcode:2017COPB...39...50R. doi:10.1016/j.pbi.2017.05.008. PMID 28601651.
  85. ^ Oldroyd GE, Harrison MJ, Paszkowski U (May 2009). "Reprogramming plant cells for endosymbiosis". Science. 324 (5928): 753–754. Bibcode:2009Sci...324..753O. doi:10.1126/science.1171644. PMID 19423817. S2CID 206518892.
  86. ^ Bianciotto V, Bandi C, Minerdi D, Sironi M, Tichy HV, Bonfante P (August 1996). "An obligately endosymbiotic mycorrhizal fungus itself harbors obligately intracellular bacteria". Applied and Environmental Microbiology. 62 (8): 3005–3010. Bibcode:1996ApEnM..62.3005B. doi:10.1128/aem.62.8.3005-3010.1996. PMC 168087. PMID 8702293.
  87. ^ Begum N, Qin C, Ahanger MA, Raza S, Khan MI, Ashraf M, et al. (2019). "Role of Arbuscular Mycorrhizal Fungi in Plant Growth Regulation: Implications in Abiotic Stress Tolerance". Frontiers in Plant Science. 10: 1068. doi:10.3389/fpls.2019.01068. PMC 6761482. PMID 31608075.
  88. ^ Herre EA, Mejía LC, Kyllo DA, Rojas E, Maynard Z, Butler A, Van Bael SA (March 2007). "Ecological implications of anti-pathogen effects of tropical fungal endophytes and mycorrhizae". Ecology. 88 (3): 550–558. Bibcode:2007Ecol...88..550H. doi:10.1890/05-1606. PMID 17503581.
  89. ^ Domka AM, Rozpaądek P, Turnau K (2019). "Are Fungal Endophytes Merely Mycorrhizal Copycats? The Role of Fungal Endophytes in the Adaptation of Plants to Metal Toxicity". Frontiers in Microbiology. 10: 371. doi:10.3389/fmicb.2019.00371. PMC 6428775. PMID 30930857.
  90. ^ Rodriguez RJ, White JF, Arnold AE, Redman RS (April 2009). "Fungal endophytes: diversity and functional roles". The New Phytologist. 182 (2): 314–330. doi:10.1111/j.1469-8137.2009.02773.x. PMID 19236579.
  91. ^ Purahong W, Hyde KD (1 March 2011). "Effects of fungal endophytes on grass and non-grass litter decomposition rates". Fungal Diversity. 47 (1): 1–7. doi:10.1007/s13225-010-0083-8. ISSN 1878-9129. S2CID 43678079.
  92. ^ "Evolutionary Development of the Clavicipitaceae". The Fungal Community: 525–538. 24 May 2005. doi:10.1201/9781420027891-33. ISBN 9780429116407.
  93. ^ Khan AL, Al-Harrasi A, Al-Rawahi A, Al-Farsi Z, Al-Mamari A, Waqas M, et al. (30 June 2016). "Endophytic Fungi from Frankincense Tree Improves Host Growth and Produces Extracellular Enzymes and Indole Acetic Acid". PLOS ONE. 11 (6): e0158207. Bibcode:2016PLoSO..1158207K. doi:10.1371/journal.pone.0158207. PMC 4928835. PMID 27359330.
  94. ^ Shahzad R, Waqas M, Khan AL, Asaf S, Khan MA, Kang SM, et al. (September 2016). "Seed-borne endophytic Bacillus amyloliquefaciens RWL-1 produces gibberellins and regulates endogenous phytohormones of Oryza sativa". Plant Physiology and Biochemistry. 106: 236–243. doi:10.1016/j.plaphy.2016.05.006. PMID 27182958.
  95. ^ Pirttilä, Anna Maria; Frank, A. Carolin (11 July 2011). Endophytes of Forest Trees: Biology and Applications. Springer Science & Business Media. ISBN 978-94-007-1599-8.
  96. ^ Senthilkumar et al., 2011
  97. ^ Quadt-Hallmann A, Kloepper JW, Benhamou N (10 February 2011). "Bacterial endophytes in cotton: mechanisms of entering the plant". Canadian Journal of Microbiology. 43 (6): 577–582. doi:10.1139/m97-081.
  98. ^ Hardoim PR, van Overbeek LS, Elsas JD (October 2008). "Properties of bacterial endophytes and their proposed role in plant growth". Trends in Microbiology. 16 (10): 463–471. doi:10.1016/j.tim.2008.07.008. PMID 18789693.
  99. ^ Bodył A, Mackiewicz P, Stiller JW (July 2007). "The intracellular cyanobacteria of Paulinella chromatophora: endosymbionts or organelles?". Trends in Microbiology. 15 (7): 295–296. doi:10.1016/j.tim.2007.05.002. PMID 17537638.
  100. ^ Asaf S, Khan MA, Khan AL, Waqas M, Shahzad R, Kim A, Kang S, Lee I (1 January 2017). "Bacterial endophytes from arid land plants regulate endogenous hormone content and promote growth in crop plants: an example of Sphingomonas sp. and Serratia marcescens". Journal of Plant Interactions. 12 (1): 31–38. Bibcode:2017JPlaI..12...31A. doi:10.1080/17429145.2016.1274060. ISSN 1742-9145. S2CID 90203067.
  101. ^ a b Jung J, Kim JS, Taffner J, Berg G, Ryu CM (1 January 2020). "Archaea, tiny helpers of land plants". Computational and Structural Biotechnology Journal. 18: 2494–2500. doi:10.1016/j.csbj.2020.09.005. PMC 7516179. PMID 33005311.
  102. ^ Taffner J, Cernava T, Erlacher A, Berg G (September 2019). "Novel insights into plant-associated archaea and their functioning in arugula (Eruca sativa Mill.)". Journal of Advanced Research. Special Issue on Plant Microbiome. 19: 39–48. doi:10.1016/j.jare.2019.04.008. PMC 6629838. PMID 31341668. S2CID 155746848.
  103. ^ Ma M, Du H, Sun T, An S, Yang G, Wang D (February 2019). "Characteristics of archaea and bacteria in rice rhizosphere along a mercury gradient". The Science of the Total Environment. 650 (Pt 1): 1640–1651. Bibcode:2019ScTEn.650.1640M. doi:10.1016/j.scitotenv.2018.07.175. PMID 30054090. S2CID 51727014.
  104. ^ Knief C, Delmotte N, Chaffron S, Stark M, Innerebner G, Wassmann R, et al. (July 2012). "Metaproteogenomic analysis of microbial communities in the phyllosphere and rhizosphere of rice". The ISME Journal. 6 (7): 1378–1390. Bibcode:2012ISMEJ...6.1378K. doi:10.1038/ismej.2011.192. PMC 3379629. PMID 22189496.
  105. ^ Moissl-Eichinger C, Pausan M, Taffner J, Berg G, Bang C, Schmitz RA (January 2018). "Archaea Are Interactive Components of Complex Microbiomes". Trends in Microbiology. 26 (1): 70–85. doi:10.1016/j.tim.2017.07.004. PMID 28826642.
  106. ^ Müller H, Berg C, Landa BB, Auerbach A, Moissl-Eichinger C, Berg G (2015). "Plant genotype-specific archaeal and bacterial endophytes but similar Bacillus antagonists colonize Mediterranean olive trees". Frontiers in Microbiology. 6: 138. doi:10.3389/fmicb.2015.00138. PMC 4347506. PMID 25784898.
  107. ^ Von Dohlen, Carol D., Shawn Kohler, Skylar T. Alsop, and William R. McManus. "Mealybug β-proteobacterial endosymbionts contain γ-proteobacterial symbionts." Nature 412, no. 6845 (2001): 433-436.
  108. ^ a b c d e f Shaffer JP, Carter ME, Spraker JE, Clark M, Smith BA, Hockett KL, et al. (April 2022). Lindemann SR (ed.). "Transcriptional Profiles of a Foliar Fungal Endophyte (Pestalotiopsis, Ascomycota) and Its Bacterial Symbiont (Luteibacter, Gammaproteobacteria) Reveal Sulfur Exchange and Growth Regulation during Early Phases of Symbiotic Interaction". mSystems. 7 (2): e0009122. doi:10.1128/msystems.00091-22. PMC 9040847. PMID 35293790.
  109. ^ Arnold AE (April 2022). "Bacterial-fungal interactions: Bacteria take up residence in the house that Fungi built". Current Biology. 32 (7): R327–R328. Bibcode:2022CBio...32.R327A. doi:10.1016/j.cub.2022.02.024. PMID 35413262. S2CID 248089525.
  110. ^ Büttner H, Niehs SP, Vandelannoote K, Cseresnyés Z, Dose B, Richter I, et al. (September 2021). "Bacterial endosymbionts protect beneficial soil fungus from nematode attack". Proceedings of the National Academy of Sciences of the United States of America. 118 (37): e2110669118. Bibcode:2021PNAS..11810669B. doi:10.1073/pnas.2110669118. PMC 8449335. PMID 34504005.
  111. ^ Villarreal LP (October 2001). "Persisting Viruses Could Play Role in Driving Host Evolution". ASM News. Archived from the original on 8 May 2009.
  112. ^ Belshaw R, Pereira V, Katzourakis A, Talbot G, Paces J, Burt A, Tristem M (April 2004). "Long-term reinfection of the human genome by endogenous retroviruses". Proceedings of the National Academy of Sciences of the United States of America. 101 (14): 4894–4899. Bibcode:2004PNAS..101.4894B. doi:10.1073/pnas.0307800101. PMC 387345. PMID 15044706.