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C Value Paradox

The C-value paradox refers to the observation that eukaryotic genome sizes vary wildly over 100,000-fold despite similar levels of complexity. Some single-celled amoebae have genomes 100 times larger than the human genome. This variation suggests genomes contain substantial non-gene DNA. The paradox is also related to the observation that the human mutation rate suggests our genome is too large to be entirely functional. Estimates in the 1970s indicated only 1-20% of human DNA may be genic, with the rest evolving neutrally. While some DNA is regulatory, the C-value paradox explained why bulk non-genic DNA amounts vary between similar species.

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Rohit Mali
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84 views

C Value Paradox

The C-value paradox refers to the observation that eukaryotic genome sizes vary wildly over 100,000-fold despite similar levels of complexity. Some single-celled amoebae have genomes 100 times larger than the human genome. This variation suggests genomes contain substantial non-gene DNA. The paradox is also related to the observation that the human mutation rate suggests our genome is too large to be entirely functional. Estimates in the 1970s indicated only 1-20% of human DNA may be genic, with the rest evolving neutrally. While some DNA is regulatory, the C-value paradox explained why bulk non-genic DNA amounts vary between similar species.

Uploaded by

Rohit Mali
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© © All Rights Reserved
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Download as DOCX, PDF, TXT or read online on Scribd
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C Value Paradox: A Summary

We might expect more complex organisms to have progressively


larger genomes, but eukaryotic genome size fails to correlate well
with apparent complexity, and instead varies wildly over more than
a 100,000-fold range. Single-celled amoebae have some of the
largest genomes, up to 100-fold larger than the human genome.
This variation suggested that genomes can contain a substantial
fraction of DNA other than for genes and their regulatory
sequences. C.A. Thomas Jr dubbed it the ‘C-value paradox’ in
1971. The C-value paradox is related to another puzzling
observation, called ‘mutational load’: the human genome seems
too large, given the observed human mutation rate. If the entire
human genome were functional (in the sense of being under
selective pressure), we would have too many deleterious mutations
per generation. By 1970, rough calculations had suggested to
several authors that maybe only 1–20% of the human genome
could be genic, with the rest evolving neutrally or nearly so. ‘C-
value’ means the ‘constant’ (or ‘characteristic’) value of haploid
DNA content per nucleus, typically measured in pictograms (1
picogram is roughly 1 gigabase). C-value paradox isn’t just an
observation that different species have different genome sizes, it’s
the observation that even similar species can have quite different
genome sizes. For example, there are many examples of related
species in the same genus that have haploid genome sizes that
differ by three- to eight fold; this is particularly common in plants,
as seen in species of rice(Oryza), Sorghum, or onions (Allium).
The maize (Zea mays) genome expanded by about 50% in just
140,000 years since its divergence from Zea luxurians (and not
merely by polyploidization). Many hypotheses have been proposed
and carefully weighed in the literature. At first, people looked for
explanations in terms of some functional significance of the extra
DNA — an adaptive function that would maintain nongenic,
nonregulatory DNA by natural selection. To explain the C-value
paradox, you have to explain why this bulk amount would vary
quite a bit even between similar species. Ohno and others in the
early 1970s, ultimately led to a reasonably well-accepted
explanation of the C-value paradox. Ohno, who believed that
strongly polarizing statements clarify scientific debate, called this
‘junk DNA’. So is all noncoding DNA, “junk DNA”? No. Of
course we’ve also known since the earliest days of molecular
biology (including the Jacob/Monod lac operon paradigm) that
genes are regulated by sequences that often occur in noncoding
DNA. Rather the idea is that there is a fraction of DNA that is
useful and functional for the organism (genes and regulatory
regions) which does more or less scale with organismal
complexity, and a ‘junk’ fraction which varies widely in amount,
creating the C-value paradox. How much nonfunctional DNA an
organism would harbor will be a tradeoff between how deleterious
it is to carry versus how easy it is to get rid of. It’s actually not
obvious that extra DNA would be all that deleterious; DNA
replication is a relatively small part of the energy budget of most
organisms. Still, DNA deletions are common enough mutations. If
there were even a small selective disadvantage to having a junky
genome, especially in species with large population sizes (where
small selection coefficient have more effect) and fast growth rates
(where an obese genome might especially be a hindrance), it would
be surprising to see a lot of nonfunctional DNA.

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