Monday, June 29, 2009


Perhaps you've thought that viruses are the simplest sort of "living" thing – if a virus can even be called "alive".

Well, maybe not. Viroids are even simpler. You're on your own as to whether you want to describe them as "alive".

Viroids are not a new discovery – they've been known since 1971. Viroids are found only in plant cells and don't seem to infect animals. They can cause plant pathology, apparently enough to be a serious economic problem.

A viroid consists entirely of a circular piece of RNA, that may be only a few hundred base units long. The smallest known viroid has only 220 units. None of this RNA codes for proteins – unlike virus RNA or DNA, which codes for proteins that (among other things) encapsulate the genetic material. The RNA or DNA of a virus is much larger than the RNA of a viroid. The smallest known virus capable of causing an infection by itself is about 2000 base units.

Viruses reproduce by co-opting machinery of the host cell. The DNA of a DNA virus, for example, is typically normal double-stranded DNA. In the virus life cycle, two separate processes are required (among others). The DNA itself has to be copied with a DNA polymerase enzyme, just as is used in making DNA copies during cell division. The proteins that the virus requires for its coat are also made in the normal way – using RNA polymerase enzyme to make messenger RNA, which can then be used to make proteins in a cell's ribosomes.

RNA viruses are trickier. Sometimes they work by using an enzyme called reverse transcriptase, which makes DNA from RNA. The HIV-1 virus responsible for AIDS is an example. Other RNA viruses, such as human polio viruses, use another enzyme, RNA replicase, which makes copies of RNA directly. RNA viruses usually encode the enzymes that they need for reproduction, to ensure a sufficient quantity of the enzyme.

So how does a viroid reproduce, given that it consists of RNA, but doesn't code for any special enzymes, or any proteins at all? The process isn't well understood, as the following explains:

Viroids: Molecular Vestiges Of The RNA World (5/17/09)
As opposed to plant viruses, which encode proteins that mediate their own replication and movement, viroids depend exclusively on host factors for these purposes. Viroids replicate through an RNA-based rolling circle mechanism with three steps: i) synthesis of longer-than-unit strands catalyzed by a host nuclear or chloroplastic RNA polymerase that reiteratively transcribes the initial circular template, ii) processing to unit-length, which remarkably is mediated by hammerhead ribozymes in the family Avsunviroidae, and iii) and circularization resulting from the action of an RNA ligase or from self-ligation.

Among the many pending issues, how viroids redirect the template specificity of certain host DNA-dependent RNA polymerases to transcribe RNA, is one of the most challenging. In addition, viroids must recruit host factors for their intracelular, cell-to-cell and long-distance movement within the plant. There are also pending questions in this context, the most appealing of which is how members of the family Avsunviridae gain access into the chloroplast; because essentially no other RNA has been reported to traffic inside this organelle, the answer to this question may reveal novel transport pathways in plant cells.

In order to understand what this is saying, the first fact needed is that viroid RNA comes in the form of a circle – no loose ends. In this respect, it is somewhat like bacterial DNA, which consists partly of circular loops of double-stranded DNA, called plasmids.

The interesting part is that viroids are apparently replicated by RNA polymerase, which normally produces RNA from a DNA template, rather than an RNA template. The process is called rolling circle replication, because the enzyme may travel around the loop a number of times, since there are no clear start and stop points. Later, in a separate operation, an RNA enzyme (ribozyme) of the host, cuts the multiply copied segments of viroid RNA back into unit segments, which join at the ends to form a circle again.

What's especially interesting about viroids is that they may give some insight into mechanisms that would be important in the RNA world hypothesis. This is the idea that before proteins existed, or even DNA itself, there was RNA – see here for some recent findings about how RNA itself may have originated.

RNA is capable of carrying genetic information just as DNA does – after all, that's what happens in RNA viruses. The main problem is how it was possible for RNA to reproduce itself. Viroids hardly give us a complete answer to this problem, since proteins (such as RNA polymerase) are still required for replication. But at least, in viroids, we have an example of a replicating entity that consists entirely of genetic information, with no proteins of its own.

Further reading:

Viroids and Virusoids

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