Preface by Elio
Bacteria, archaea, microfungi, and many protists are "mainstream" in the sense that everyone in our business is aware of them. Other highly consequential microbes such as the microsporidia and the oomycetes commonly elicit reactions such as "I really should look into them sometime." I have been a microbiologist for upward of sixty years but have paid only the scantest attention to these large and important groups of organisms (and therefore hang my head in shame). But it's never too late to correct such oversights. We are starting out by reproducing, with permission of the author, a particularly illuminating article on the microsporidia by Patrick Keeling.
by Patrick Keeling
What Are Microsporidia?
Microsporidia are a diverse group of obligate intracellular eukaryotic parasites. There are approximately 1,300 formally described species in 160 genera [1], but this certainly represents a tiny fraction of the real diversity because most potential host lineages have been poorly surveyed. Nearly all microsporidia are known to infect animals, and some are responsible for a number of human diseases (13 species of microsporidia have been documented to infect humans) predominantly associated with immune suppression [2]. They also infect several commercially important animal species such as bees, silk worms, and salmon, and various domesticated mammals. They are thought to be especially common in insects and fish, although most invertebrates have been so poorly surveyed this may change.
Figure 1. Light micrograph of Antonospora locustae with pressure-
induced polar tube eversion. The scale bar is 10 µm. (A) Many
ungerminated spores (one example labeled U) and a few germinated
spores, showing the residual spore wall (one example labeled G).
(B) A germinated spore where the everted polar tube (PT) has
extended far from the cell and can be seen to be many times the
length of the spore.
Their infective stage is a thick-walled spore, which is also the only stage that can survive outside their host cell [3]. The spore contains a sophisticated infection apparatus, primarily distinguished by a long, coiled filament called the polar filament. When the spore germinates, an inflow of water leads to pressure in the spore that eventually ruptures the wall and forces the polar filament to eject, turning inside out to form a tube (Figure 1) [4]. This process takes place very quickly, so the polar tube is in effect a projectile. At the completion of germination, the parasite cytoplasm is forced through the tube and either delivered to the surface of the host cell, or perhaps injected into the host cytoplasm if the projectile tube has actually penetrated the host cell. It has also been shown that microsporidia can be taken up by phagocytosis, and then use the polar tube to escape from the vacuole [5], so there appear to be more than one mode of infection.
Are They Protists, Fungi, or What?
There has been considerable debate about the origin of microsporidian parasites. Aside from their elaborate infection mechanism, they have few distinguishing features, and have thus been difficult to compare to other eukaryotes. This is illustrated by their tumultuous taxonomic history (Figure 2), and the tendency to lump them with what we now know to be unrelated organisms. When microsporidia were discovered in 1857, they were considered to be schizomycete fungi, but this was an artificial group that included various yeasts and bacteria. They were soon transferred to Sporozoa and ultimately to the subgroup Cnidosporidia. This too was a grab-bag of four unrelated groups (Microsporidia, Myxosporidia, Actinosporidia, and Helicosporidia) that were falsely grouped because of their intracellular parasitic way of life. Remarkably, Microsporidia, Myxosporidia, and Helicosporidia have since been shown to be fungi, animals, and green algae, respectively, underscoring just how distantly related these parasites really are.