Endosymbiont
An endosymbiont or endobiont
Endosymbiosis played key roles in the development of
Symbionts are either obligate (require their host to survive) or facultative (can survive independently).
Etymology
Endosymbiosis comes from the Greek: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living".
Symbiogenesis
Typically the endosymbiont's genome shrinks, discarding genes whose roles are displaced by the host.
Transmission
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
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
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]
Hosts
Invertebrates
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]
Insects
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]
Primary
Among primary endosymbionts of insects, the best-studied are the pea
In some insect groups, these endosymbionts live in specialized insect cells called
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
Primary endosymbionts of insects have among the smallest of known bacterial genomes and have
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
Secondary
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
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]
Marine
Extracellular endosymbionts are represented in all four extant classes of
Some marine
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]
Dinoflagellates
Previously thought to be a single species, molecular
Phytoplankton
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
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
Protists
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]
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
Vertebrates
The spotted salamander (
Plants
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
Fungal endophytes can be found in all plant tissues. Fungi living below the ground amidst plant roots are known as
Arbuscular Mycorrhizal Fungi (AMF)
Endophytic fungi
Endophytic fungi in
Endophytic bacteria
Endophytic bacteria belong to a diverse group of plant endosymbionts characterized by systematic colonization of plant tissues. The most common genera include
Archaea endosymbionts
The characterization of archaea includes crop plants such as
Bacteria
Some Betaproteobacteria have Gammaproteobacteria endosymbionts.[107]
Fungi
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
Virus endosymbionts
The
See also
- Epibiont, organism living on the surface of another organism
- Anagenesis
- Endophyte
- Ectosymbiosis
- List of symbiotic organisms
- List of symbiotic relationships
- Multigenomic organism
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