Endosymbiont - Knowledge

588: 406: 1605:. 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. Most plant endosymbiosis studies focus on fungal or bacteria using 216: 289:) is a process where a host acquires a facultative symbiont from the environment or another host. The Rhizobia-Legume symbiosis (bacteria-plant endosymbiosis) is a prime example of this modality. 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 33: 1020:. There are some asymbiotic (occurs without an endosymbiont) Rhizosolenia, however there appears to be mechanisms limiting growth of these organisms in low nutrient conditions. 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. 535:

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'

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genome is generally intact. While other species like that of the UNCY-A symbiont and Richelia have reduced genomes. 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

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signals that the legume detects, leading to root nodule formation. This process bleeds into other processes such as nitrogen fixation in plants. The evolutionary advantage of such an interaction allows genetic exchange between both organisms involved to increase the propensity for novel functions as

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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

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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

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living as endosymbionts in plants can enhance their host's primary productivity either by producing or capturing important resources. These endosymbionts can also enhance plant productivity by producing toxic metabolites that aid plant defenses against

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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".

664:" that attack the molecular machinery of invading parasites. 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. 314:

Vertical transmission takes place when the symbiont moves directly from parent to offspring. In horizontal transmission each generation acquires symbionts from the environment. An example is nitrogen-fixing bacteria in certain plant roots, such as

230:. 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. 6125:

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".

1413:, in planta colonization and host plant type. Clavicipitaceous fungi systematically colonize temperate season grasses. Non-clavicipitaceous fungi colonize higher plants and even roots and divide into subcategories. 672:

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

3247:"Figure 5—figure supplement 2. KEGG metabolic reconstructions based on the intact genes present in the Acromyrmex, Solenopsis, Apis mellifera and Anopheles gambiae genomes, together constituting the urea cycle". 543:

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

471:. 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. 1327:, 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 901:. 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 3947:"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" 978:

well above intracellular requirements, and found the cyanobacterium was likely fixing nitrogen for its host. Additionally, both host and symbiont cell growth were much greater than free-living

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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.

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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.

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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.

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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

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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

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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.

1194:. 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 5848:

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".

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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".

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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. Fungal

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concentrations. Endosymbiotic bacteria fix nitrogen for their hosts and in turn receive organic carbon from photosynthesis. These symbioses play an important role in global

1148:. 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 6971: 1385:

lives as a plant endosymbiont and also harbors further endosymbiont intracytoplasmic bacterium-like organisms. AMF generally promote plant health and growth and alleviate

6312:"Bacterial endophytes from arid land plants regulate endogenous hormone content and promote growth in crop plants: an example of Sphingomonas sp. and Serratia marcescens" 521:

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.

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Fungi host endohyphal bacteria; the effects of the bacteria are not well studied. Many such fungi in turn live within plants. These fungi are otherwise known as fungal

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Aksoy, S., Pourhosseini, A. & Chow, A. 1995. Mycetome endosymbionts of tsetse flies constitute a distinct lineage related to Enterobacteriaceae. Insect Mol Biol.

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

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organelles. The chloroplast is derived from a cyanobacterial primary endosymbiosis that began over one billion years ago. An oxygenic, photosynthetic free-living

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Symbionts are either obligate (require their host to survive) or facultative (can survive independently). The most common examples of obligate endosymbiosis are

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Endophytic bacteria belong to a diverse group of plant endosymbionts characterized by systematic colonization of plant tissues. The most common genera include

2257: 554:. Studying insect endosymbionts can aid understanding the origins of symbioses in general, as a proxy for understanding endosymbiosis in other species. 1393:. Individual AMF species have different effects in different hosts – introducing the AMF of one plant to another plant can reduce the latter's growth. 1315:

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

767:) have obligate extracellular endosymbionts that fill the entire body of their host. These marine worms are nutritionally dependent on their symbiotic 319:

symbionts. A third type is mixed-mode transmission, where symbionts move horizontally for some generations, after which they are acquired vertically.

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Aksoy S, Maudlin I, Dale C, Robinson AS, O'Neill SL (January 2001). "Prospects for control of African trypanosomiasis by tsetse vector manipulation".

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deleterious mutations and small mutations from the population, resulting in a loss of genes over many millions of years. Research in which a parallel

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Jaenike J, Unckless R, Cockburn SN, Boelio LM, Perlman SJ (July 2010). "Adaptation via symbiosis: recent spread of a Drosophila defensive symbiont".

482:), and are maternally-transmitted, i.e. the mother transmits her endosymbionts to her offspring. In some cases, the bacteria are transmitted in the 1371: 368:. In the event of a bottleneck, a decrease in symbiont diversity could compromise host-symbiont interactions, as deleterious mutations accumulate. 6588:

Moissl-Eichinger C, Pausan M, Taffner J, Berg G, Bang C, Schmitz RA (January 2018). "Archaea Are Interactive Components of Complex Microbiomes".

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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".

6030: 747:. Other studies indicate that these subcuticular bacteria may be both abundant within their hosts and widely distributed among the Echinoderms. 1545:

family. Endophytic bacteria typically colonize the leaf tissues from plant roots, but can also enter the plant through the leaves through leaf

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nitrogen independently. This endosymbiont reduction in genome size, might be a step that occurred in the evolution of organelles (above).

4283:"Influence of the Amazon River plume on distributions of free-living and symbiotic cyanobacteria in the western tropical north Atlantic Ocean" 4055:"Chloroplast genes are expressed during intracellular symbiotic association of Vaucheria litorea plastids with the sea slug Elysia chlorotica" 132:

that produce energy from sunlight. Some 100 million years ago, UCYN-A, a nitrogen-fixing bacterium, became an endosymbiont of the marine alga

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Jeon KW, Jeon MS (October 1976). "Endosymbiosis in amoebae: recently established endosymbionts have become required cytoplasmic components".

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Bodył A, Mackiewicz P, Stiller JW (July 2007). "The intracellular cyanobacteria of Paulinella chromatophora: endosymbionts or organelles?".

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Scientists classify insect endosymbionts as Primary or Secondary. Primary endosymbionts (P-endosymbionts) have been associated with their

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with their insect hosts. Secondary endosymbionts more recently associated with their hosts, may be horizontally transferred, live in the

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Baker AC (November 2003). "Flexibility and Specificity in Coral-Algal Symbiosis: Diversity, Ecology, and Biogeography of Symbiodinium".

1632:. This archaeal abundance is associated with plant species type, environment and the plant’s developmental stage. In a study on plant 494:, they are transmitted via milk to the embryo. In termites, the endosymbionts reside within the hindguts and are transmitted through 348:

symbionts have lost genes for essential molecules and rely on the host to supply them. In return, the symbionts synthesize essential

1246:. Endophytes aid in processes such as growth and development, nutrient uptake, and defense against biotic and abiotic stresses like 3204:

Douglas AE (January 1998). "Nutritional interactions in insect-microbial symbioses: aphids and their symbiotic bacteria Buchnera".

4829: 4547:"Seasonal dynamics of the endosymbiotic, nitrogen-fixing cyanobacterium Richelia intracellularis in the eastern Mediterranean Sea" 528:

and competition from other bacterial species, the ample supply of nutrients and relative environmental stability inside the host.

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a tsetse fly symbiont, is vertically transmitted (via mother's milk). When a symbiont reaches this stage, it resembles a cellular

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Smith, Noel H.; Gordon, Stephen V.; de la Rua-Domenech, Ricardo; Clifton-Hadley, Richard S.; Hewinson, R. Glyn (September 2006).

5893:"Are Fungal Endophytes Merely Mycorrhizal Copycats? The Role of Fungal Endophytes in the Adaptation of Plants to Metal Toxicity" 5084: 6625:"Plant genotype-specific archaeal and bacterial endophytes but similar Bacillus antagonists colonize Mediterranean olive trees" 3797:"Generality of toxins in defensive symbiosis: Ribosome-inactivating proteins and defense against parasitic wasps in Drosophila" 3746:"Generality of toxins in defensive symbiosis: Ribosome-inactivating proteins and defense against parasitic wasps in Drosophila" 2016: 459: 7794: 6180: 5260:"The Hidden World within Plants: Ecological and Evolutionary Considerations for Defining Functioning of Microbial Endophytes" 5121: 5025: 4973: 4946: 4936: 4839: 4197:

Villareal T (1994). "Widespread occurrence of the Hemiaulus-cyanobacterial symbiosis in the southwest North Atlantic Ocean".

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have two bacterial endosymbionts. Ruthmannia lives inside the animal's digestive cells. Grellia lives permanently inside the

397:(insects). Many insect agricultural pests and human disease vectors have intimate relationships with primary endosymbionts. 6872: 5447:"Playing Chutes and Ladders: Heterogeneity and the Relative Roles of Bottom-Up and Top-Down Forces in Natural Communities" 1636:-specific archaeal and bacterial endophytes, 35% of archaeal sequences were detected in overall sequences (achieved using 7799: 5136:

K. Jeon, “Amoeba and X-bacteria: Symbiont Acquisition and Possible Species Change,” in: L. Margulis and R. Fester, eds.,

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events while True endophytes possess adaptive traits because of which they live strictly in association with plants. The

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of the plant tissue in a sterile environment. Passenger endophytic bacteria eventually colonize inner tissue of plant by

6068:"Endophytic Fungi from Frankincense Tree Improves Host Growth and Produces Extracellular Enzymes and Indole Acetic Acid" 3291:"Origin of Mutualism Between Termites and Flagellated Gut Protists: Transition From Horizontal to Vertical Transmission" 2838: 2384: 1620:, but also aquatic plants. The abundance of archaea varies by tissue type; for example archaea are more abundant in the 727:). Little is known of the nature of the association (mode of infection, transmission, metabolic requirements, etc.) but 1069:

that lacks mitochondria. However, spherical bacteria live inside the cell and serve the function of the mitochondria.

6050: 2875:"Transmission of Bacterial Symbionts With and Without Genome Erosion Between a Beetle Host and the Plant Environment" 623:

defends its aphid host from parasitoid wasps. This symbiosis replaces lost elements of the insect's immune response.

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of bacteria and insects was inferred supports the belief that primary endosymbionts are transferred only vertically.

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hosts for millions of years (from ten to several hundred million years). They form obligate associations and display

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Typically the endosymbiont's genome shrinks, discarding genes whose roles are displaced by the host. For example the

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Welburn SC, Maudlin I, Ellis DS (June 1987). "In vitro cultivation of rickettsia-like-organisms from Glossina spp".

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Little AF, van Oppen MJ, Willis BL (June 2004). "Flexibility in algal endosymbioses shapes growth in reef corals".

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Fungal endophytes can be found in all plant tissues. Fungi living below the ground amidst plant roots are known as

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have photosynthetic symbionts. In some species the host digests algae to keep the population at a constant level.

1832: 344:, the symbionts do not need to survive independently, often leading them to have a reduced genome. For instance, 6694:

Shaffer JP, Carter ME, Spraker JE, Clark M, Smith BA, Hockett KL, et al. (April 2022). Lindemann SR (ed.).

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and enzyme production within its host, which, in turn, may influence the effect the fungus has on its plant host

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Porras-Alfaro A, Bayman P (8 September 2011). "Hidden fungi, emergent properties: endophytes and microbiomes".

2620:"Infection and Invasion of Roots by Symbiotic, Nitrogen-Fixing Rhizobia during Nodulation of Temperate Legumes" 1852: 5410:

Schardl CL, Leuchtmann A, Spiering MJ (2 June 2004). "Symbioses of grasses with seedborne fungal endophytes".

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or nutrient-poor regions of the ocean like that of the North Atlantic. In such waters, cell growth of larger

17: 7409: 7090: 5799:"Role of Arbuscular Mycorrhizal Fungi in Plant Growth Regulation: Implications in Abiotic Stress Tolerance" 1550: 7095: 1903:

Mergaert P (April 2018). "Role of antimicrobial peptides in controlling symbiotic bacterial populations".

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Zchori-Fein E, Perlman SJ (July 2004). "Distribution of the bacterial symbiont Cardinium in arthropods".

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is a marine ciliate that uses endosymbiotic, purple, non-sulphur bacteria for anoxygenic photosynthesis.

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Hardoim PR, van Overbeek LS, Berg G, Pirttilä AM, Compant S, Campisano A, et al. (September 2015).

250:, allowing their genomes to diversify. Selection is episodic (when the cicadas reproduce). The original 4661:"Division cycles in the nitrogen-fixingRhizosolenia(Bacillariophyceae)-Richelia(Nostocaceae) symbiosis" 1533: 1523: 4739: 7630: 6812:

Büttner H, Niehs SP, Vandelannoote K, Cseresnyés Z, Dose B, Richter I, et al. (September 2021).

2076:"Genomic divergence within non-photosynthetic cyanobacterial endosymbionts in rhopalodiacean diatoms" 1181:

the algae is transferred to only one of the daughter cells, while the other cell restarts the cycle.

445: 134: 4740:"Identification of the ectosymbiotic bacteria of Mixotricha paradoxa involved in movement symbiosis" 7477: 7378: 7295: 1781: 1698:

These interactions may impact the way that fungi interact with the environment by modulating their

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Khan AL, Al-Harrasi A, Al-Rawahi A, Al-Farsi Z, Al-Mamari A, Waqas M, et al. (30 June 2016).

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Subramaniam A, Yager PL, Carpenter EJ, Mahaffey C, Björkman K, Cooley S, et al. (July 2008).

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genome split into three much simpler endosymbionts, each encoding only a few genes—an instance of

7504: 7280: 6533:"Metaproteogenomic analysis of microbial communities in the phyllosphere and rhizosphere of rice" 5742:"An obligately endosymbiotic mycorrhizal fungus itself harbors obligately intracellular bacteria" 5689:

Oldroyd GE, Harrison MJ, Paszkowski U (May 2009). "Reprogramming plant cells for endosymbiosis".

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Roth R, Paszkowski U (October 2017). "Plant carbon nourishment of arbuscular mycorrhizal fungi".

4332:"Amazon River enhances diazotrophy and carbon sequestration in the tropical North Atlantic Ocean" 4225:"Extensive bloom of a N2-fixing diatom/cyanobacterial association in the tropical Atlantic Ocean" 551: 286: 267: 4545:

Zeev EB, Yogev T, Man-Aharonovich D, Kress N, Herut B, Béjà O, Berman-Frank I (September 2008).

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Klein A, Schrader L, Gil R, Manzano-Marín A, Flórez L, Wheeler D, et al. (February 2016).

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that lives within the body or cells of another organism. Typically the two organisms are in a

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Knief C, Delmotte N, Chaffron S, Stark M, Innerebner G, Wassmann R, et al. (July 2012).

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Carpenter EJ, Montoya JP, Burns J, Mulholland MR, Subramaniam A, Capone DG (20 August 1999).

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Primary endosymbionts of insects have among the smallest of known bacterial genomes and have

271: 7068: 6708:) Reveal Sulfur Exchange and Growth Regulation during Early Phases of Symbiotic Interaction" 4601:"Genomic deletions disrupt nitrogen metabolism pathways of a cyanobacterial diatom symbiont" 7436: 7431: 7307: 6907: 6825: 6768: 6544: 6489: 6323: 6135: 6079: 5857: 5753: 5698: 5655: 5458: 5363:"Fungal plant endosymbionts alter life history and reproductive success of aphid predators" 5163: 4612: 4558: 4507: 4450: 4396: 4294: 4236: 4114:"Deceptively simple: Minute marine animals live in a sophisticated symbiosis with bacteria" 4066: 4011: 3958: 3903: 3698: 3643: 3488: 3394: 2693: 2567: 2431: 2204: 2087: 2028: 1947: 1773: 1743:

Another interesting example of a bacterium living in symbiosis with a fungus is the fungus

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with a complicated feeding apparatus that feeds on other microbes. When it engulfs a green

668: 609: 603: 506: 167: 37: 2941: 2806: 2789: 2352: 2335: 8: 7672: 7514: 7470: 7426: 7164: 6757:"Bacterial-fungal interactions: Bacteria take up residence in the house that Fungi built" 5510:"Algal endosymbionts as vectors of horizontal gene transfer in photosynthetic eukaryotes" 3477:"A novel intracellular mutualistic bacterium in the invasive ant Cardiocondyla obscurior" 2511:

Wernegreen JJ (November 2002). "Genome evolution in bacterial endosymbionts of insects".

2017:"Scientists discover first algae that can fix nitrogen — thanks to a tiny cell structure" 1676: 1637: 1409:

are taxonomically diverse and are divided into categories based on mode of transmission,

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for the aphid host. When a symbiont reaches this stage, it begins to resemble a cellular

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Belshaw R, Pereira V, Katzourakis A, Talbot G, Paces J, Burt A, Tristem M (April 2004).

6829: 6772: 6548: 6493: 6327: 6139: 6083: 5861: 5757: 5702: 5659: 5599:"Endosymbionts in cranberry: Diversity, effect on plant growth, and pathogen biocontrol" 5572: 5462: 5167: 4616: 4562: 4511: 4454: 4400: 4298: 4240: 4070: 4015: 3962: 3907: 3702: 3647: 3492: 3398: 2925: 2697: 2571: 2478: 2435: 2285:"An Expanded Ribosomal Phylogeny of Cyanobacteria Supports a Deep Placement of Plastids" 2208: 2091: 2032: 1951: 1354:

endosymbionts as endo and ecto as well. AMF plant endosymbionts systematically colonize

405: 7789: 7722: 6848: 6813: 6794: 6732: 6695: 6651: 6624: 6565: 6532: 6513: 6462: 6449: 6420: 6396: 6371: 6349: 6102: 6067: 6012: 5919: 5892: 5825: 5798: 5722: 5628: 5536: 5509: 5490: 5482: 5387: 5362: 5335: 5308: 5284: 5259: 5235: 5210: 5186: 5151: 5066: 4996: 4917: 4912: 4895: 4805: 4778: 4720: 4633: 4600: 4496:"Diatom fluxes to the deep sea in the oligotrophic North Pacific gyre at Station Aloha" 4471: 4438: 4358: 4331: 4312: 4170: 4035: 3927: 3823: 3796: 3772: 3745: 3721: 3686: 3667: 3611: 3584: 3560: 3535: 3511: 3476: 3229: 3162: 3127: 3108: 3048: 3023: 2999: 2974: 2901: 2874: 2819: 2765: 2730: 2729:

Chrostek, Ewa; Pelz-Stelinski, Kirsten; Hurst, Gregory D. D.; Hughes, Grant L. (2017).

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or AMF are the most diverse plant microbial endosymbionts. With exceptions such as the

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This soil-dwelling fungus lives in close association with a toxin-producing bacteria,

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on coral reefs and its role in coral bleaching is an important in coral reef ecology.

177:, thrive in their intermediate insect hosts because of an obligate endosymbiosis with 7717: 7421: 6935: 6853: 6798: 6786: 6737: 6656: 6605: 6570: 6505: 6466: 6454: 6401: 6341: 6310:

Asaf S, Khan MA, Khan AL, Waqas M, Shahzad R, Kim A, Kang S, Lee I (1 January 2017).

6292: 6257: 6176: 6151: 6107: 6046: 6004: 5965: 5960: 5943: 5924: 5873: 5830: 5779: 5726: 5714: 5671: 5632: 5620: 5576: 5541: 5474: 5427: 5392: 5340: 5289: 5240: 5191: 5117: 5088: 5058: 5021: 4969: 4942: 4876: 4857:"An endosymbiotic Chlorella-bearing ciliate: Platyophrya chlorelligera Kawakami 1989" 4835: 4810: 4759: 4724: 4712: 4638: 4576: 4476: 4412: 4408: 4363: 4281:

Foster RA, Subramaniam A, Mahaffey C, Carpenter EJ, Capone DG, Zehr JP (March 2007).

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All vascular plants harbor endosymbionts or endophytes in this context. They include

806: 790: 780: 247: 6517: 6501: 6353: 6016: 5765: 5494: 5309:"Multifaceted Interactions Between Endophytes and Plant: Developments and Prospects" 5070: 5000: 4921: 4316: 4174: 3970: 3931: 3233: 3186: 2556:"Idiosyncratic Genome Degradation in a Bacterial Endosymbiont of Periodical Cicadas" 2540: 1975: 1549:. Generally, the endophytic bacteria are isolated from the plant tissues by surface 1474:

and abnormal reproduction, suggesting that the mycotoxins are transmitted along the

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spp., and has a reduced genome. A 2011 study measured nitrogen fixation by the

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has spread across North America owing to its ability to defend its fly host against

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In some insect groups, these endosymbionts live in specialized insect cells called

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of the insects (not specialized bacteriocytes, see below), and are not obligate.

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spp. They can both be eliminated by treatments that target their bacterial host.

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