How do the products of the mitochondrial and plastid genomes contribute to the f

unction of these organelles?

Both mitochondria and plastids were originally bacteria which formed an endosymb
iotic relationship with eukaryotes. They had the full bacterial genomes needed
to produce all the proteins they needed to survive. But over time the genes for
many necessary components have been passed to the nucleus.
Now the mitochondrial genome only contains only the genes for ribosomal RNAs, tr
ansfer RNAs, ribosomal proteins and proteins for the electron transfer chain and
ATP synthesis. In some species with larger mitochondrial genomes, the genome a
lso encodes some genes for membrane protein assembly. We can see this process o
f gene transfer to the nucleus in the legumes. The Cox II gene has been transfe
rred to the nucleus in one lineage and this has allowed mitochondrial gene loss
in one line of its descendants.
Nuclear genes encode more than 90% of organelle proteins, but the organelle geno
mes still play an important role. Both mitochondria and plastids are able to co
ntrol nuclear gene expression. Adding tagetitoxin, an inhibitor of chloroplast
RNA polymerase prevents nuclear gene expression earlier in seedling development,
demonstrating that the plastid genome is able to control nuclear transcription.
The exact mechanism by which the signal is passed is not known.
We see how important the mitochondrial genome is when it becomes mutated. Mild
missense mutatons can give rise to diseases such as Leber s hereditary optic neuro
pathy, since it results in less electron transfer and ATP production which are c
rucial for neurons. This can result in the loss of sight at an age between 20 a
nd 24. In other cases point deletions occur which would be lethal but for the f
act that there still remain some functional mitochondria (heteroplasmy). The mi
tochondria is a dangerous environment for DNA due to reactive oxygen species, wh
ich is one driving force towards moving genes to the nucleus. Mitochondrial DNA
mutations have been shown to have a role in aging both in brains and muscles of
people over 80.
Chloroplast genomes tend to have an inverted repeat which maintains their stabi
lity. Hybridisation has shown that as with mitochondria they encode tRNas and r
RNAs. This is crucial since it is not believed that RNAs can be imported into e
ither the mitochondria or the plastids.
While we typically think of a plastid s chief function as performing photosynthesi
s but in fact they perform other functions and are found in non photosynthetic s
pecies such as Plasmodium and Toxoplasma. These plastids with short 35kbp geno
mes are known as apicoplasts, they perform protein synthesis and are essential f
or the cells that contain them. The essential functions of their genomes mean t
hat the antibiotic lincomycin can be used as an antimalarial.
In a number of cases where a complex is formed, some subunits are encoded by gen
es in the chloroplast and others by genes in the nucleus. In such cases the oth
er units are translated in the cytoplasm and then imported into the plastid. Th
is allows control by both components. Chloroplast genomes encode the large subu
nit of rubisco and components of both photosystems, cytochrome bf and ATP syntha
se. These are all crucial to the function of a chloroplast.
Chloroplasts contain two enzymes for polymerisation of RNA, plastid-encoded poly
merase (PEP) and nuclear-encoded polymerase (NEP). As the names would apply PEP
genes are found in the chloroplast DNA. PEP is used to transcribe genes for ph
otosynthesis. There is also a nuclear encoded polymerase (NEP) which is similar
to bacteriophage polymerases. It is transcribed and translated outside the pla
stid. It is imported and transcribes the chloroplast DNA for genetic systems.
Non-green plastids (amyloplasts and leucoplasts) are found in the roots of photo
synthetic plants. They are still transcribing the same genes but levels of poly
merase are lower than in the leaves. This allows a degree of control over the ty
pe of plastid. While there are no activators or repressors at the level of tran
scription there is translational regulation and post translational regulation by
the rapid degradation of subunits which have not been assembled into a complex.
This means that if the nucleus is not producing a subunit a complex will not b
e able to be formed.
The proteins found in both plastids and mitochondria come from a combination of
the nuclear genome and the organelle genome. Movement to the nucleus has afford
ed protection for DNA but the organelle genome is still crucial to allow the syn
thesis of RNAs and the localised control of protein expression.