Endosymbiont
Encyclopedia
An endosymbiont is any organism
that lives within the body or cells of another organism, i.e. forming an endosymbiosis (Greek
: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living"). Examples are nitrogen-fixing bacteria
(called rhizobia
) which live in root nodules on legume roots, single-celled algae
inside reef-building corals, and bacterial endosymbionts that provide essential nutrients to about 10%–15% of insects.
Many instances of endosymbiosis are obligate- that is, either the endosymbiont or the host cannot survive without the other, such as the gutless marine worms
of the genus
Riftia, which get nutrition from their endosymbiotic bacteria. The most common examples of obligate endosymbiosis are mitochondria and chloroplasts. Some human parasites, e.g. : Wucherichia bancrofti and Mansonella perstans thrive in their hosts because of an obligate endosymbiosis with Wolbachi spp.. They can both be eliminated from their host by treatments that target this bacterium. However, not all endosymbioses are obligate. Also, some endosymbioses can be harmful
to either of the organisms involved.
It is generally agreed that certain organelle
s of the eukaryotic
cell, especially mitochondria and plastid
s such as chloroplast
s, originated as bacterial endosymbionts. This theory is called the endosymbiotic theory
, and was first articulated by the Russia
n botanist Konstantin Mereschkowski
in 1905.
According to endosymbiosis theory, an anaerobic cell probably ingested an aerobic
bacterium but failed to digest it. The aerobic bacterium flourished within the cell because the cell's cytoplasm
was abundant in half-digested food molecule
s. The bacterium digested these molecules with oxygen
and gained great amounts of energy. Because the bacterium had so much energy, it probably leaked some of it as Adenosine triphosphate
into the cell's cytoplasm. This benefited the anaerobic cell because it enabled it to digest food aerobically. Eventually, the aerobic bacterium could no longer live independently from the cell, and it therefore became a mitochondrion. The origin of the chloroplast
is very similar to that of the mitochondrion. A cell must have captured a photosynthetic
cyanobacterium and failed to digest it. The cyanobacterium thrived in the cell and eventually evolved into the first chloroplast. Other eukaryotic organelles may have also evolved through endosymbiosis; it has been proposed that cilia, flagella, centriole
s, and microtubule
s may have originated from a symbiosis between a Spirochaete
bacterium and an early eukaryotic cell, but this is not widely accepted among biologists.
There are several examples of evidence that support endosymbiosis theory. Mitochondria and chloroplasts contain their own small supply of DNA
, which may be remnants of the genome
the organelles had when they were independent aerobic bacteria. The single most convincing evidence of the descent of organelles from bacteria is the position of mitochondria and plastid DNA sequences in phylogenetic tree
s of bacteria. Mitochondria have sequences that clearly indicate origin from a group of bacteria called the alpha-Proteobacteria. Plastids have DNA sequences that indicate origin from the cyanobacteria (blue-green algae). In addition, there are organisms alive today, called living intermediates, that are in a similar endosymbiotic condition to the prokaryotic cells and the aerobic bacteria. Living intermediates show that the evolution proposed by the endosymbiont theory is possible. For example, the giant amoeba Pelomyxa
lacks mitochondria but has aerobic bacteria that carry out a similar role. A variety of coral
s, clam
s, snail
s, and one species of Paramecium
permanently host algae
in their cells. Many of the insect endosymbionts have been shown to have ancient associations with their hosts, involving strictly vertical inheritance. In addition, these insect symbionts have similar patterns of genome evolution
to those found in true organelles: genome reduction, rapid rates of gene evolution, and bias in nucleotide
base composition favoring adenine
and thymine
, at the expense of guanine
and cytosine
.
Further evidence of endosymbiosis are the prokaryotic ribosomes found within chloroplasts and mitochondria as well as the double-membrane enclosing them. It used to be widely assumed that the inner membrane is the original membrane of the once independent prokaryote, while the outer one is the food vacuole (phagosomal membrane) it was enclosed in initially. However, this view neglects the fact that i) both modern cyanobacteria and alpha-proteobacteria are Gram negative bacteria, which are surrounded by double membranes; ii) the outer membranes of the endosymbiotic organelles (chloroplasts and mitochondria) are very similar to those of these bacteria in their lipid and protein compositions. Accumulating biochemical data strongly suggest that the double membrane enclosing chloroplasts and mitochondria derived from those of the ancestral bacteria, and the phagosomal membrane disappeared during organelle evolution. Triple or quadruple membranes are found among certain algae, probably resulting from repeated endosymbiosis (although little else was retained of the engulfed cell).
These modern organisms with endosymbiotic relationships with aerobic bacteria have verified the endosymbiotic theory, which explains the origin of mitochondria and chloroplasts from bacteria. Researchers in molecular and evolutionary biology no longer question this theory, although some of the details, such as the mechanisms for loss of gene
s from organelles to host nuclear genomes, are still being worked out.
, 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 in general.
Some marine oligochaeta
(e.g. Olavius or Inanidrillus) 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).
endosymbionts of the genus Symbiodinium, commonly known as zooxanthella
e, are found in corals, mollusks (esp. giant clam
s, the Tridacna), sponges
, and foraminifera
. These endosymbionts drive the amazing formation of coral reefs by capturing sunlight and providing their hosts with energy for carbonate
deposition.
Previously thought to be a single species, molecular phylogenetic evidence over the past couple decades has shown there to be great diversity in Symbiodinium. In some cases there is specificity between host and Symbiodinium clade. More often, however, there is an ecological distribution of Symbiodinium, the symbionts switching between hosts with apparent ease. When reefs become environmentally stressed, this distribution of symbionts 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 presents one of the most complex and interesting current problems in reef ecology
.
is a protozoan that lacks mitochondria, however, spherical bacteria live inside the cell and serve the function of the mitochondria. Mixotricha also has three other species of symbionts that live on the surface of the cell.
Paramecium bursaria
, a species of ciliate
, has a mutualistic symbiotic relationship with green alga called Zoochlorella. The algae live inside the cell, in the cytoplasm.
hosts for many millions of years (from 10 to several hundred million years in some cases), they form obligate associations (see below), and display cospeciation with their insect hosts. Secondary endosymbionts exhibit a more recently developed association, are sometimes horizontally transferred between hosts, live in the hemolymph
of the insects (not specialized bacteriocytes, see below), and are not obligate.
Among primary endosymbionts of insects, the best studied are the pea aphid
(Acyrthosiphon pisum) and its endosymbiont Buchnera
sp. APS, the tsetse fly
Glossina morsitans morsitans and its endosymbiont Wigglesworthia glossinidia brevipalpis
and the endosymbiotic protists in lower termite
s. As with endosymbiosis in other insects, the symbiosis is obligate in that neither the bacteria nor the insect is viable without the other. Scientists have been unable to cultivate the bacteria in lab conditions outside of the insect. With special nutritionally-enhanced diets, the insects can survive, but are unhealthy, and at best survive only a few generations.
In some insect groups, these endosymbionts live in specialized insect cells called bacteriocyte
s (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 developing insect embryo. In termites, the endosymbionts reside within the hindguts and are transmitted through trophallaxis
among colony members.
The primary endosymbionts are thought to help the host either by providing nutrients that the host cannot obtain itself, or by metabolizing insect waste products into safer forms. For example, the putative primary role of Buchnera is to synthesize essential amino acid
s that the aphid cannot acquire from its natural diet of plant sap. Similarly, the primary role of Wigglesworthia is probably to synthesize vitamin
s 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 which constitutes 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.
Genome sequencing reveals that obligate bacterial endosymbionts of insects have among the smallest of known bacterial genomes and have lost many genes that are commonly found in closely related bacteria. Several theories have been put forth to explain the loss of genes. Presumably 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 the primary endosymbionts are transferred only vertically (i.e. from the mother), and not horizontally (i.e. by escaping the host and entering a new host).
Attacking obligate bacterial endosymbionts may present a way to control their insect hosts, many of which are pests or carriers of human disease. For example aphids are crop pests and the tsetse fly carries the organism Trypanosoma brucei
that causes African sleeping sickness. Other motivations for their study is to understand symbiosis, and to understand how bacteria with severely depleted genomes are able to survive, thus improving our knowledge of genetics
and molecular biology
.
Less is known about secondary endosymbionts. The pea aphid (Acyrthosiphon pisum) is known to contain at least three secondary endosymbionts, Hamiltonella defensa, Regiella insecticola, and Serratia symbiotica. H. defensa aids in defending the insect from parasitoids. Sodalis glossinidius is a secondary endosymbiont tsetse flies that lives inter- and intracellularly in various host tissues, including the midgut and hemolymph. Phylogenetic studies have not indicated a correlation between evolution of Sodalis and tsetse. Unlike tsetse's P-symbiont Wigglesworthia, though, Sodalis has been cultured in vitro.
in order to limit the exchange of migratory cells between the developing embryo and the body of the mother, where an epithelium
won't do because certain blood cells are specialized to be able to insert themselves between adjacent epithelial cells. The ERV is a virus similar to HIV
(the virus causing AIDS
in humans). The immunodepressive action was the initial normal behavior of the virus, similar to HIV. The fusion proteins was a way to spread the infection to other cells by simply merging them with the infected one (similar to HIV). It is believed that the ancestors of modern vivipary
mammals evolved after an accidental infection of an ancestor with this virus, that permitted the fetus to survive the immune system of the mother.
The human genome project found several thousand ERVs, which are organized into 24 families.
Organism
In biology, an organism is any contiguous living system . In at least some form, all organisms are capable of response to stimuli, reproduction, growth and development, and maintenance of homoeostasis as a stable whole.An organism may either be unicellular or, as in the case of humans, comprise...
that lives within the body or cells of another organism, i.e. forming an endosymbiosis (Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living"). Examples are nitrogen-fixing bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
(called rhizobia
Rhizobia
Rhizobia are soil bacteria that fix nitrogen after becoming established inside root nodules of legumes . Rhizobia require a plant host; they cannot independently fix nitrogen...
) which live in root nodules on legume roots, single-celled algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...
inside reef-building corals, and bacterial endosymbionts that provide essential nutrients to about 10%–15% of insects.
Many instances of endosymbiosis are obligate- that is, either the endosymbiont or the host cannot survive without the other, such as the gutless marine worms
Siboglinidae
Siboglinidae, also known as the beard worms, is a family of polychaete annelid worms whose members made up the former phyla Pogonophora and Vestimentifera. They are composed of about 100 species of vermiform creatures and live in thin tubes buried in sediments at ocean depths from 100 to 10,000 m...
of the genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Riftia, which get nutrition from their endosymbiotic bacteria. The most common examples of obligate endosymbiosis are mitochondria and chloroplasts. Some human parasites, e.g. : Wucherichia bancrofti and Mansonella perstans thrive in their hosts because of an obligate endosymbiosis with Wolbachi spp.. They can both be eliminated from their host by treatments that target this bacterium. However, not all endosymbioses are obligate. Also, some endosymbioses can be harmful
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...
to either of the organisms involved.
It is generally agreed that certain organelle
Organelle
In cell biology, an organelle is a specialized subunit within a cell that has a specific function, and is usually separately enclosed within its own lipid bilayer....
s of the eukaryotic
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...
cell, especially mitochondria and plastid
Plastid
Plastids are major organelles found in the cells of plants and algae. Plastids are the site of manufacture and storage of important chemical compounds used by the cell...
s such as chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...
s, originated as bacterial endosymbionts. This theory is called the endosymbiotic theory
Endosymbiotic theory
The endosymbiotic theory concerns the mitochondria, plastids , and possibly other organelles of eukaryotic cells. According to this theory, certain organelles originated as free-living bacteria that were taken inside another cell as endosymbionts...
, and was first articulated by the Russia
Russia
Russia or , officially known as both Russia and the Russian Federation , is a country in northern Eurasia. It is a federal semi-presidential republic, comprising 83 federal subjects...
n botanist Konstantin Mereschkowski
Konstantin Mereschkowski
Konstantin Mereschcowsky was a prominent Russian biologist, botanist and advocate of eugenics active mainly around Kazan, whose research on lichens led him to propose the theory of symbiogenesis - that larger, more complex cells evolved from the symbiotic relationship between less complex ones...
in 1905.
Endosymbiosis theory and mitochondria and chloroplasts
The endosymbiosis theory attempts to explain the origins of organelles such as mitochondria and chloroplasts in eukaryotic cells. The theory proposes that chloroplasts and mitochondria evolved from certain types of bacteria that eukaryotic cells engulfed through endophagocytosis. These cells and the bacteria trapped inside them entered a symbiotic relationship, a close association between different types of organisms over an extended time. However, more specifically, the relationship was endosymbiotic, meaning that one of the organisms (the bacteria) lived within the other (the eukaryotic cells).According to endosymbiosis theory, an anaerobic cell probably ingested an aerobic
Aerobic organism
An aerobic organism or aerobe is an organism that can survive and grow in an oxygenated environment.Faculitative anaerobes grow and survive in an oxygenated environment and so do aerotolerant anaerobes.-Glucose:...
bacterium but failed to digest it. The aerobic bacterium flourished within the cell because the cell's cytoplasm
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...
was abundant in half-digested food molecule
Molecule
A molecule is an electrically neutral group of at least two atoms held together by covalent chemical bonds. Molecules are distinguished from ions by their electrical charge...
s. The bacterium digested these molecules with oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
and gained great amounts of energy. Because the bacterium had so much energy, it probably leaked some of it as Adenosine triphosphate
Adenosine triphosphate
Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...
into the cell's cytoplasm. This benefited the anaerobic cell because it enabled it to digest food aerobically. Eventually, the aerobic bacterium could no longer live independently from the cell, and it therefore became a mitochondrion. The origin of the chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...
is very similar to that of the mitochondrion. A cell must have captured a photosynthetic
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...
cyanobacterium and failed to digest it. The cyanobacterium thrived in the cell and eventually evolved into the first chloroplast. Other eukaryotic organelles may have also evolved through endosymbiosis; it has been proposed that cilia, flagella, centriole
Centriole
A Centriole is a barrel-shaped cell structure found in most animal eukaryotic cells, though it is absent in higher plants and most fungi. The walls of each centriole are usually composed of nine triplets of microtubules...
s, and microtubule
Microtubule
Microtubules are a component of the cytoskeleton. These rope-like polymers of tubulin can grow as long as 25 micrometers and are highly dynamic. The outer diameter of microtubule is about 25 nm. Microtubules are important for maintaining cell structure, providing platforms for intracellular...
s may have originated from a symbiosis between a Spirochaete
Spirochaete
Spirochaetes belong to a phylum of distinctive Gram-negative bacteria, which have long, helically coiled cells...
bacterium and an early eukaryotic cell, but this is not widely accepted among biologists.
There are several examples of evidence that support endosymbiosis theory. Mitochondria and chloroplasts contain their own small supply of DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
, which may be remnants of the genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
the organelles had when they were independent aerobic bacteria. The single most convincing evidence of the descent of organelles from bacteria is the position of mitochondria and plastid DNA sequences in phylogenetic tree
Tree
A tree is a perennial woody plant. It is most often defined as a woody plant that has many secondary branches supported clear of the ground on a single main stem or trunk with clear apical dominance. A minimum height specification at maturity is cited by some authors, varying from 3 m to...
s of bacteria. Mitochondria have sequences that clearly indicate origin from a group of bacteria called the alpha-Proteobacteria. Plastids have DNA sequences that indicate origin from the cyanobacteria (blue-green algae). In addition, there are organisms alive today, called living intermediates, that are in a similar endosymbiotic condition to the prokaryotic cells and the aerobic bacteria. Living intermediates show that the evolution proposed by the endosymbiont theory is possible. For example, the giant amoeba Pelomyxa
Pelomyxa
Pelomyxa are giant amoebae, usually 500-800 μm but occasionally up to 5 mm in length. One notable species is P. palustris; other described species may be synonyms, or have been moved to the unrelated genus Chaos...
lacks mitochondria but has aerobic bacteria that carry out a similar role. A variety of coral
Coral
Corals are marine animals in class Anthozoa of phylum Cnidaria typically living in compact colonies of many identical individual "polyps". The group includes the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "head" is a colony of...
s, clam
Clam
The word "clam" can be applied to freshwater mussels, and other freshwater bivalves, as well as marine bivalves.In the United States, "clam" can be used in several different ways: one, as a general term covering all bivalve molluscs...
s, snail
Snail
Snail is a common name applied to most of the members of the molluscan class Gastropoda that have coiled shells in the adult stage. When the word is used in its most general sense, it includes sea snails, land snails and freshwater snails. The word snail without any qualifier is however more often...
s, and one species of Paramecium
Paramecium
Paramecium is a group of unicellular ciliate protozoa, which are commonly studied as a representative of the ciliate group, and range from about 0.05 to 0.35 mm in length. Simple cilia cover the body, which allow the cell to move with a synchronous motion at speeds of approximately 12 body...
permanently host algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...
in their cells. Many of the insect endosymbionts have been shown to have ancient associations with their hosts, involving strictly vertical inheritance. In addition, these insect symbionts have similar patterns of genome evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
to those found in true organelles: genome reduction, rapid rates of gene evolution, and bias in nucleotide
Nucleotide
Nucleotides are molecules that, when joined together, make up the structural units of RNA and DNA. In addition, nucleotides participate in cellular signaling , and are incorporated into important cofactors of enzymatic reactions...
base composition favoring adenine
Adenine
Adenine is a nucleobase with a variety of roles in biochemistry including cellular respiration, in the form of both the energy-rich adenosine triphosphate and the cofactors nicotinamide adenine dinucleotide and flavin adenine dinucleotide , and protein synthesis, as a chemical component of DNA...
and thymine
Thymine
Thymine is one of the four nucleobases in the nucleic acid of DNA that are represented by the letters G–C–A–T. The others are adenine, guanine, and cytosine. Thymine is also known as 5-methyluracil, a pyrimidine nucleobase. As the name suggests, thymine may be derived by methylation of uracil at...
, at the expense of guanine
Guanine
Guanine is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine . In DNA, guanine is paired with cytosine. With the formula C5H5N5O, guanine is a derivative of purine, consisting of a fused pyrimidine-imidazole ring system with...
and cytosine
Cytosine
Cytosine is one of the four main bases found in DNA and RNA, along with adenine, guanine, and thymine . It is a pyrimidine derivative, with a heterocyclic aromatic ring and two substituents attached . The nucleoside of cytosine is cytidine...
.
Further evidence of endosymbiosis are the prokaryotic ribosomes found within chloroplasts and mitochondria as well as the double-membrane enclosing them. It used to be widely assumed that the inner membrane is the original membrane of the once independent prokaryote, while the outer one is the food vacuole (phagosomal membrane) it was enclosed in initially. However, this view neglects the fact that i) both modern cyanobacteria and alpha-proteobacteria are Gram negative bacteria, which are surrounded by double membranes; ii) the outer membranes of the endosymbiotic organelles (chloroplasts and mitochondria) are very similar to those of these bacteria in their lipid and protein compositions. Accumulating biochemical data strongly suggest that the double membrane enclosing chloroplasts and mitochondria derived from those of the ancestral bacteria, and the phagosomal membrane disappeared during organelle evolution. Triple or quadruple membranes are found among certain algae, probably resulting from repeated endosymbiosis (although little else was retained of the engulfed cell).
These modern organisms with endosymbiotic relationships with aerobic bacteria have verified the endosymbiotic theory, which explains the origin of mitochondria and chloroplasts from bacteria. Researchers in molecular and evolutionary biology no longer question this theory, although some of the details, such as the mechanisms for loss of gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...
s from organelles to host nuclear genomes, are still being worked out.
Bacterial endosymbionts in marine invertebrates
Extracellular endosymbionts are also 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 alpha group of the class ProteobacteriaProteobacteria
The Proteobacteria are a major group of bacteria. They include a wide variety of pathogens, such as Escherichia, Salmonella, Vibrio, Helicobacter, and many other notable genera....
, 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 in general.
Some marine oligochaeta
Oligochaeta
Oligochaeta is a subclass of animals in the biological phylum Annelida, which is made up of many types of aquatic and terrestrial worms, and this includes all of the various earthworms...
(e.g. Olavius or Inanidrillus) 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).
Symbiodinium dinoflagellate endosymbionts in marine metazoa and protists
DinoflagellateDinoflagellate
The dinoflagellates are a large group of flagellate protists. Most are marine plankton, but they are common in fresh water habitats as well. Their populations are distributed depending on temperature, salinity, or depth...
endosymbionts of the genus Symbiodinium, commonly known as zooxanthella
Zooxanthella
Zooxanthellae are flagellate protozoa that are golden-brown intracellular endosymbionts of various marine animals and protozoa, especially anthozoans such as the scleractinian corals and the tropical sea anemone, Aiptasia....
e, are found in corals, mollusks (esp. giant clam
Giant clam
The giant clam, Tridacna gigas , is the largest living bivalve mollusc. T. gigas is one of the most endangered clam species. It was mentioned as early as 1825 in scientific reports...
s, the Tridacna), sponges
Sea sponge
Sponges are animals of the phylum Porifera . Their bodies consist of jelly-like mesohyl sandwiched between two thin layers of cells. While all animals have unspecialized cells that can transform into specialized cells, sponges are unique in having some specialized cells, but can also have...
, and foraminifera
Foraminifera
The Foraminifera , or forams for short, are a large group of amoeboid protists which are among the commonest plankton species. They have reticulating pseudopods, fine strands of cytoplasm that branch and merge to form a dynamic net...
. These endosymbionts drive the amazing formation of coral reefs by capturing sunlight and providing their hosts with energy for carbonate
Carbonate
In chemistry, a carbonate is a salt of carbonic acid, characterized by the presence of the carbonate ion, . The name may also mean an ester of carbonic acid, an organic compound containing the carbonate group C2....
deposition.
Previously thought to be a single species, molecular phylogenetic evidence over the past couple decades has shown there to be great diversity in Symbiodinium. In some cases there is specificity between host and Symbiodinium clade. More often, however, there is an ecological distribution of Symbiodinium, the symbionts switching between hosts with apparent ease. When reefs become environmentally stressed, this distribution of symbionts is related to the observed pattern of coral bleaching
Coral bleaching
Coral bleaching is the loss of intracellular endosymbionts through either expulsion or loss of algal pigmentation.The corals that form the structure of the great reef ecosystems of tropical seas depend upon a symbiotic relationship with unicellular flagellate protozoa, called zooxanthellae, that...
and recovery. Thus the distribution of Symbiodinium on coral reefs and its role in coral bleaching presents one of the most complex and interesting current problems in reef ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...
.
Endosymbionts in protists
Mixotricha paradoxaMixotricha paradoxa
Mixotricha paradoxa is a species of protozoan that lives inside the termite species Mastotermes darwiniensis and has multiple bacterial symbionts. The name, given by the Australian biologist J.L. Sutherland, who first described Mixotricha in 1933,. means “the paradoxical being with mixed-up...
is a protozoan that lacks mitochondria, however, spherical bacteria live inside the cell and serve the function of the mitochondria. Mixotricha also has three other species of symbionts that live on the surface of the cell.
Paramecium bursaria
Paramecium bursaria
Paramecium bursaria is a species of ciliate protozoan that has a mutualistic symbiotic relationship with green algae called Zoochlorella. The algae live inside the Paramecium in its cytoplasm and provide it with food, while the Paramecium provides the algae with movement and protection. P...
, a species of ciliate
Ciliate
The ciliates are a group of protozoans characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagella but typically shorter and present in much larger numbers with a different undulating pattern than flagella...
, has a mutualistic symbiotic relationship with green alga called Zoochlorella. The algae live inside the cell, in the cytoplasm.
Bacterial endosymbionts in insects
Scientists classify insect endosymbionts in two broad categories, 'Primary' and 'Secondary'. Primary endosymbionts (sometimes referred to as P-endosymbionts) have been associated with their insectInsect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
hosts for many millions of years (from 10 to several hundred million years in some cases), they form obligate associations (see below), and display cospeciation with their insect hosts. Secondary endosymbionts exhibit a more recently developed association, are sometimes horizontally transferred between hosts, live in the hemolymph
Hemolymph
Hemolymph, or haemolymph, is a fluid in the circulatory system of some arthropods and is analogous to the fluids and cells making up both blood and interstitial fluid in vertebrates such as birds and mammals...
of the insects (not specialized bacteriocytes, see below), and are not obligate.
Among primary endosymbionts of insects, the best studied are the pea aphid
Aphid
Aphids, also known as plant lice and in Britain and the Commonwealth as greenflies, blackflies or whiteflies, are small sap sucking insects, and members of the superfamily Aphidoidea. Aphids are among the most destructive insect pests on cultivated plants in temperate regions...
(Acyrthosiphon pisum) and its endosymbiont Buchnera
Buchnera (proteobacteria)
Buchnera aphidicola a member of the Proteobacteria, is the primary endosymbiont of aphids and has been studied in the pea aphid, Acyrthosiphon pisum. It is believed that Buchnera was once a free living gram negative ancestor similar to a modern Enterobacteriaceae such as Escherichia coli...
sp. APS, the tsetse fly
Tsetse fly
Tsetse , sometimes spelled tzetze and also known as tik-tik flies, are large biting flies that inhabit much of mid-continental Africa between the Sahara and the Kalahari deserts. They live by feeding on the blood of vertebrate animals and are the primary biological vectors of trypanosomes, which...
Glossina morsitans morsitans and its endosymbiont Wigglesworthia glossinidia brevipalpis
Wigglesworthia glossinidia brevipalpis
Wigglesworthia glossinidia is a Gram-negative bacterium in the family Enterobacteriaceae, related to E. coli, which lives in the gut of the tsetse fly. The bacterium was described by Serap Aksoy and bears the name of the British entomologist Sir Vincent Brian Wigglesworth who died the year prior...
and the endosymbiotic protists in lower termite
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
s. As with endosymbiosis in other insects, the symbiosis is obligate in that neither the bacteria nor the insect is viable without the other. Scientists have been unable to cultivate the bacteria in lab conditions outside of the insect. With special nutritionally-enhanced diets, the insects can survive, but are unhealthy, and at best survive only a few generations.
In some insect groups, these endosymbionts live in specialized insect cells called bacteriocyte
Bacteriocyte
A bacteriocyte , also called a mycetocyte, is a specialized adipocyte found in some insect groups such as aphids, tsetse flies, german cockroaches, and many others. Mycetocyte symbionts specifically include three insect groups: Cockroaches, Homoptera such as aphid, and Coleoptera such as weevils...
s (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
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
, as in Buchnera; in others like Wigglesworthia, they are transmitted via milk to the developing insect embryo. In termites, the endosymbionts reside within the hindguts and are transmitted through trophallaxis
Trophallaxis
Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth or anus-to-mouth feeding. It is most highly developed in social insects such as ants, termites, wasps and bees. The word was introduced by the entomologist William Morton Wheeler in 1918...
among colony members.
The primary endosymbionts are thought to help the host either by providing nutrients that the host cannot obtain itself, or by metabolizing insect waste products into safer forms. For example, the putative primary role of Buchnera is to synthesize essential amino acid
Essential amino acid
An essential amino acid or indispensable amino acid is an amino acid that cannot be synthesized de novo by the organism , and therefore must be supplied in the diet.-Essentiality vs. conditional essentiality in humans:...
s that the aphid cannot acquire from its natural diet of plant sap. Similarly, the primary role of Wigglesworthia is probably to synthesize vitamin
Vitamin
A vitamin is an organic compound required as a nutrient in tiny amounts by an organism. In other words, an organic chemical compound is called a vitamin when it cannot be synthesized in sufficient quantities by an organism, and must be obtained from the diet. Thus, the term is conditional both on...
s that the tsetse fly does not get from the blood
Blood
Blood is a specialized bodily fluid in animals that delivers necessary substances such as nutrients and oxygen to the cells and transports metabolic waste products away from those same cells....
that it eats. In lower termites, the endosymbiotic protists play a major role in the digestion of lignocellulosic materials which constitutes 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.
Genome sequencing reveals that obligate bacterial endosymbionts of insects have among the smallest of known bacterial genomes and have lost many genes that are commonly found in closely related bacteria. Several theories have been put forth to explain the loss of genes. Presumably 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 the primary endosymbionts are transferred only vertically (i.e. from the mother), and not horizontally (i.e. by escaping the host and entering a new host).
Attacking obligate bacterial endosymbionts may present a way to control their insect hosts, many of which are pests or carriers of human disease. For example aphids are crop pests and the tsetse fly carries the organism Trypanosoma brucei
Trypanosoma brucei
Trypanosoma brucei is a parasitic protist species that causes African trypanosomiasis in humans and nagana in animals in Africa. There are 3 sub-species of T. brucei: T. b. brucei, T. b. gambiense and T. b. rhodesiense.These obligate parasites have two hosts - an insect vector and mammalian host...
that causes African sleeping sickness. Other motivations for their study is to understand symbiosis, and to understand how bacteria with severely depleted genomes are able to survive, thus improving our knowledge of genetics
Genetics
Genetics , a discipline of biology, is the science of genes, heredity, and variation in living organisms....
and molecular biology
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...
.
Less is known about secondary endosymbionts. The pea aphid (Acyrthosiphon pisum) is known to contain at least three secondary endosymbionts, Hamiltonella defensa, Regiella insecticola, and Serratia symbiotica. H. defensa aids in defending the insect from parasitoids. Sodalis glossinidius is a secondary endosymbiont tsetse flies that lives inter- and intracellularly in various host tissues, including the midgut and hemolymph. Phylogenetic studies have not indicated a correlation between evolution of Sodalis and tsetse. Unlike tsetse's P-symbiont Wigglesworthia, though, Sodalis has been cultured in vitro.
Viral endosymbionts and endogenous retrovirus
During pregnancy in viviparous mammals, endogenous retrovirii (ERVs) are activated and produced in high quantities during the implantation of the embryo. On one hand they act as immunodepressors, and protect the embryo from the immune system of the mother and on the other hand viral fusion proteins cause the formation of the placental syncytiumSyncytium
In biology, a syncytium is a large cell-like structure; filled with cytoplasm and containing many nuclei. Most cells in eukaryotic organisms have a single nucleus; syncytia are specialized forms used by various organisms.The term may also refer to cells that are connected by specialized membrane...
in order to limit the exchange of migratory cells between the developing embryo and the body of the mother, where an epithelium
Epithelium
Epithelium is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. Epithelial tissues line the cavities and surfaces of structures throughout the body, and also form many glands. Functions of epithelial cells include secretion, selective...
won't do because certain blood cells are specialized to be able to insert themselves between adjacent epithelial cells. The ERV is a virus similar to HIV
HIV
Human immunodeficiency virus is a lentivirus that causes acquired immunodeficiency syndrome , a condition in humans in which progressive failure of the immune system allows life-threatening opportunistic infections and cancers to thrive...
(the virus causing AIDS
AIDS
Acquired immune deficiency syndrome or acquired immunodeficiency syndrome is a disease of the human immune system caused by the human immunodeficiency virus...
in humans). The immunodepressive action was the initial normal behavior of the virus, similar to HIV. The fusion proteins was a way to spread the infection to other cells by simply merging them with the infected one (similar to HIV). It is believed that the ancestors of modern vivipary
Vivipary
Vivipary has two different meanings. In animals, it means development of the embryo inside the body of the mother, eventually leading to live birth, as opposed to laying eggs...
mammals evolved after an accidental infection of an ancestor with this virus, that permitted the fetus to survive the immune system of the mother.
The human genome project found several thousand ERVs, which are organized into 24 families.
Obligate bacterial endosymbionts in insects
- A general review of bacterial endosymbionts in insects. P. Baumann, N. A. Moran and L. Baumann, Bacteriocyte-associated endosymbionts of insects in M. Dworkin, ed., The prokaryotes, Springer, New York, 2000. http://link.springer.de/link/service/books/10125/