Insects and microorganisms symbiosis: the endosymbionts

Symbiotic relationships are an important motor for organisms’ diversification and evolution. The relationships insects have established with some endosymbiotic microorganisms (that is, those inhabiting the inner of their bodies) have provided them of a lot of surprising physiological and ecological adaptations. 

The value of the relationship between insects and their endosymbionts

The major cause for insects’ evolutive and adaptive success is their potential to stablish beneficial relationships with other life beings and, especially, with those microorganisms inhabiting their insides: the endosymbionts.

Some years ago, it was considered that the greatest contribution of endosymbiotic microorganisms to the physiology of insects was their role in feeding habits, which would explain, at least in part, the diversity of diets among insects. However, it has been shown that endosymbionts affect many other physiological traits.

Types of endosymbiosis in insects

Endosymbiotic microorganisms can be found inside the gut, in the spaces between cells and inside cells.

Generally, the more internal the endosymbiotic microorganisms are within the host’s body, the closer their relationship with the insect is. The four most common types of endosymbiosis in insects are explained below, from the most external and least close relationship to the most internal and closest one.

Gut microbes

Gut microbiota of insects is composed both of prokaryotes (unicellular, without nucleus, like bacteria and archaea) and eukaryotes (unicellular or pluricellular, with nucleus, like protozoans) that live outside the gut cells. They usually inhabit the hind part of insect’s gut (hindgut), either moving freely in its lumen or remaining attached to its walls. In some phytophagous insects, likes termites and cockroaches, the hindgut is a chamber without oxygen (anaerobic) where fermentation of cellulose and other complex sugars takes place.

 

Worker termite gut; the green part corresponds to the hindgut without oxygen. Figure belonging to the following paper: Brune, A. (2014). Symbiotic digestion of lignocellulose in termite guts. Nature Reviews Microbiology, 12(3), 168-180.

In termites, this anaerobic chamber contains facultative anaerobic prokaryotes (they can develop either with or without oxygen) and obligate anaerobic prokaryotes (they can only develop without oxygen), such as spirochetes and methanogens, which aid in digestion. In addition, in some worker termites, this chamber also contains protozoans that play a major role in the digestion of wood cellulose (Have you ever seen a piece of furniture pierced by termites?).

Unlike other endosymbionts, gut microbes are horizontally transmitted between insects; that is, insects don’t inherit gut microbes from their parents, but they should acquire them throughout their lives. In termites, acquisition of gut microbes takes place through a process called trophallaxis: the workers, which are the only able to feed by themselves, digest the food and transmit the resulting product mixed with gut microorganisms to the rest of the colony members through their mouthparts.

Trophollaxis. Picture by Shutterstock.

Moreover, microorganisms are removed during molting processes, so termites (and other insects performing trophollaxis) can acquire them again through trophollaxis.

Endoparasites

Parasites that live and/or develop inside an organism are known as endoparasites. They are also horizontally transmitted between insects.

Insects stablish fairly more relationships with pluricellular endoparasites than with microorganisms, being the pluricellular endoparasites the most harmful for insects in general terms; these are the cases of insect parasitoids (of which we talked in this post) and nematodes (able to transmit deathful bacteria to insects).

The most relevant endoparasitic relationship between insects and microorganisms, and the only one we are going to explain here, are vectors: the insect (or vector) serve as a container to the parasite until it reaches the definitive host. Parasites transported by vector usually are pathogenic protozoans harmful to vertebrates, like Trypanosoma (Chagas disease), Leishmania (leishmaniosis) or Plasmodium (Malaria).

Mosquito of the genus Anopheles, the major vector of the protozoan causing malaria worldwide: Plasmodium. Public domain image.

Extracellular and intracellular symbiosis

Unlike gut microbes and endoparasites, extracellular and intracellular endosymbionts are vertically transmitted generation after generation; that is, the insect inherits them from its parents

  • Extracellular endosymbionts

Extracellular endosymbionts, which can be both prokaryotes and eukaryotes, can be found in different organs of the body (even in the intestine along with the gut microbes). In any case, they never penetrate inside the cells. However, some species can be found outside and inside cells.

Since many extracellular microorganisms can also be intracellular, the possibility that they are found, in an evolutionary sense, in a transition stage between gut microbes and intracellular endosymbionts has been discussed.

An interesting case of extracellular endosymbiosis takes place in some species of aphids of the tribe Cerataphidini. Generally, aphids stablish a close relationship with an intracellular endosymbiont bacteria (Buchnera), but in some species of the aforementioned tribe these bacteria are substituted by extracellular unicellular yeast-like fungi (YLS or ‘yeast-like symbiont’) which inhabit the cavities between organs and inside different adipose bodies. Like Buchnera in the rest of aphids, YLS would play a key role on aphid feeding habits, participating in the production of essential nutrients.

Ceratovacuna nekoashi (Cerataphidini). Link (CC 2.5)

It is suggested that YLS would have evolved from an entomopathogenic fungus (that is, harmful to insects) whose lineage would later have derived into beneficial endosymbiotic organisms.

  • Intracellular endosymbionts

It is considered that at least 70% of insects has endosymbiotic microorganisms inside its cells. There exist two types of intracellular endosymbionts:

Mycetocyte symbionts or Blochmann bodies

Bacteriocytes or mycetocytes are specialized adipose cells containing endosymbionts which can be found in some groups of insects. These cells are vertically transmitted to the offspring and gathered together forming organs known as mycetomes o bacteriomes.

Blochmann bodies, or simply the endosymbionts inside mycetomes, are related to three groups of insects: Blattaria (cockroaches), some groups of heteropterans within Homoptera (cicadas, rust flies, aphids, etc.) and Curculionidae (curculionid beetles).

Buchnera aphidicola inside a mycetome of the aphid Acyrthosiphon pisum. The central element is the mycetome’s nucleus. Buchnera cells, which are round, are located packed in the citoplasm of the mycetome. Picture by J. White y N. Moran, University of Arizona (CC 2.5).

The most well studied case is the relationship between Buchnera and aphids. This intracellular bacterium recycles the uric acid and some other nitrogenous wastes produced by the aphid in order to produce the amino acid glutamine, which is then used by this same endosymbiont to produce other essential amino acids necessary for the aphid to develop. It is also considered that Buchnera produces vitamin B2 (riboflavin). This can explain why aphids have such a high reproductive rate and a big evolutive success despite having a diet rich in carbohydrates (which they obtain from plant’s sap) and poor in nitrogenous compounds.

It has been confirmed that Buchnera cells decrease in number when nutrients are scarce. This suggests that aphids use Buchnera cells as an alternative food source in difficult situations. So, aphids take more advantages from this relationship than Buchnera.

Guest endosymbionts

In this case, the guest (endosymbiont) alters some physiological traits of the insect to obtain some advantage.

Guest endosymbionts usually affect the sex ratio of insects (proportion of males and females in a population) as well as other reproductive traits. Guest endosymbionts that alter the sex ratio are known as sex-ratio distorters. Some guest microbes inhabiting the cytoplasm of insect’s cells are vertically transmitted to the offspring through ovules, so they need a higher proportion of female insects to guarantee their own perpetuity. To alter this proportion, they use different methods: male killing, induction of parthenogenesis, feminization or cytoplasm incompatibility, for which they usually induce changes at the genetic level.

One of the most well-studied cases is Wolbachia, an intracellular bacterium capable to induce a sex-ratio bias through almost every of the aforementioned methods.

Phenotypes resulting from insects infected with Wolbachia. Figure belonging to the following paper: Werren, J. H., Baldo, L. & Clark, M. E. 2008. Wolbachia: master manipulators of invertebrate biology. Nature Reviews Microbiology, 6(10), 741-751.

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Do you know any other relationship between microbes and insects? Leave your comments below!

References

  • Bourtzis K. Miller T. A. (2003). Insect Symbiosis. CRC Press.
  • Douglas, A.E. (1998). Nutritional interactions in insect-microbial symbioses: Aphids and their symbiotic bacteria Buchnera. Annual Review of Entomology, 43: 17–38.
  • Vega F.E., Blackwell M. (2005). Insect-Fungal Associations: Ecology and Evolution. Oxford University Press, USA.

The cover image is a montage made by the author from two images: 1) bacterium vector (by Flaticon from www.flaticon.com) and 2) termite vector (obtained from www.allstatepest.com.au).

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