Monday, September 25, 2006

X Chromosome Inactivation


That cute kitten is Copycat or CC – the first cloned cat. She is genetically identical to her “mom” (to the right) but phenotypically different. The reason is because of the peculiar phenomenon called X-chromosome inactivation (XCI) that occurs in all the female cells in mammals.
In mammals sex is determined by differential inheritance of the sex chromosomes. Females are XX while males are XY. To compensate for having an extra dosage of X chromosome genes that might be developmentally fatal, the cells in the female embryo randomly select one X chromosome and shut it off (transcriptionally silence it). This “silent” state is maintained throughout life in the female cell. Since the phenomenon is random (each cell independently decides its fate), mammalian females are essentially mosaics. CC is different from her progenitor because the gene for coat color lies on the X chromosome and is randomly silenced in the clone, giving it it’s unique calico pattern.
The phenomenon is evolutionarily conserved in the mammalian lineage – since there is preliminary evidence to suggest that XCI occurs in Prototherians (egg laying mammals) who arose 200-300 million years ago and also in Metatherains (marsupials). Since X and Y differentiation also occurs in these lineages, it is likely that XCI arose as a way of dosage compensation.
How is it done? The random X chromosome inactivation has been linked to a region on the X chromosome, called Xic (X- inactivation center). What is so unique about this region is that all the genes characterized so far are non- coding genes – they make the RNA but this RNA is never used to make a cellular protein. The ncRNA (non coding RNA) from this region are responsible for counting the number of X chromosomes present in the cell, marking one for silencing and maintaining the silenced state. The most talked about transcript from this region is Xist which is essential for the silencing step.
The mechanism of how the ncRNA form Xist or other mapped locus do the inactivation is not yet known. It is most likely that transcription of these genes help to bring in chromatin remodeling factors (things that signal that the chromosome is silenced). Since Xist transcription would take about 30 minutes, during which the RNA is physically attached to the X chromosome, this explanation seems plausible. The unraveling of the mechanism would help us understand another dosage compensation mechanism (that of imprinting) but that is another blog.

50 comments:

Prashanth said...

Nice one :) I find it interesting.

Psssst... change the label... put it under a broader category like Genetics or something.

One thing I did not understand in this... what exactly do you mean by the extra dosage of X chromosome being developmentally fatal? Is this related to reinforcement of genetic defects?

Lalit Singh said...

x supression
and then they say men are supressing women
abe nature hi supress kar raha hain
and we get the blame

Sakshi said...

@ Prashanth - Thanks. I will edit the label to Genetics.
The Y chromosome in humans is stripped down to ~ 80 genes whereas the X has abt 1500 odd genes. While most of the Y chromosome genes are involved in directing the development in male specific manner (testis development and such), the X chromosome has genes in addition to those of the female development. So a male embryo gets 1X dose of the genes and a female embryo would potentially get 2X dose of these genes.
Very few genes are deleterious if present in only one copy (haploinsufficiency) or are overtly deleterious in three copies. Having more or fewer copies (aneuploidy) of a large fraction of the genome is, however, invariably incompatible with viability.
And hence it is necessary for the organism to balance this dosage effect.
So a way to equalize the gene doses is to either increase the amount made from the singel X copy (fruit fly operates such) or decrease the dose from the 2Xs.
I hope that addresses the question.

Sakshi said...

@ Lalit :P
I look at it as females being so utterlyunique that not even two cells in our bodies are alike.

Prashanth said...

Actually, you lost me, but I'll take your word for it :). I'm not much of a biologist, after all...

The_Girl_From_Ipanema said...

cool!!! and i felt very much at home. :) nice sakshi!

Born a said...

I did not know about XCI. Very cool article. Like Prashanth, didnt exactly get why the dosage of the genes have to be balanced. I mean what goes wrong developmentally if one of the copies of X-chromosome is not suppressed. I know of several genes in microbes which are present in duplicate copies. Do you think that one or more copies of the gene are suppressed in that case too?

Sakshi said...

@BaL and Prashanth -
THink of it as each male and female needs X amount of a protein from each gene. Now for males that is the default amount being produced since they have just 1 chromosome. But the females has 2 copies of X and hence twice the amt of protein present in the female cell is going to be more or not equal to that of the male. Hence to keep the level of gene dosage on X chromosome same, the cells inactivate one X.
Also, duplication of chromosome is going to be fatal to the developing embryo (just a duplication of a chr. 21 leads to down's sndrome.. here we are talking abt a huge X xhromosome with few thousand genes).
The reasoning, for why dosage compensation is done, in terms of Biochemistry was actually put into a hypothesis in a neat series of papers in 1948 by Muller (I will try and find the papers and put up a link.
Does that clarify the problem?

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HEre is a link to Muller's papers I was talking about along with a commentary -
http://post.queensu.ca/~forsdyke/xchromos.htm

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