Carotenoids
are organic pigments that play contrasting roles during photosynthesis in
absorbing light energy and protecting plants from excess of light. Roberto
Bassi’s research group now reveals that carotenoids have yet a new trick up
their sleeve.
Plants and other photosynthetic
organisms live in a catch-22 situation. “Plants produce oxygen but are also
poisoned by oxygen,” says Roberto Bassi, an Italian plant physiologist who has
been passionate about photosynthesis since his graduate degree at the Padua
University Botanical Garden. Bassi’s research group at Verona University played
a pivotal role in understanding the dual function of carotenoid pigments in
absorbing light energy and protecting the photosynthetic machinery against light-induced
damage by oxygen. Now his team has identified a new unexpected function for
carotenoids in controlling the production of photosynthetic proteins.
Carotenoids are organic
pigments made by plants, algae, fungi and cyanobacteria that are found in all
organisms (animals obtain them from food). Besides their fundamental roles in
photosynthesis, carotenoids can act as plant hormones, vitamins, odours, colours
(in fruits, flowers and bird feathers, for instance) and, in the eye retina, photo-protection. There are two types of carotenoids: carotenes, which give
carrots their orange colour, and their oxygenated offshoots, the yellow
xanthophylls. In plant leaves, carotenoids are normally masked by chlorophyll,
but they put on a show in autumn as chlorophyll gets degraded and their
striking orange and yellow colours are revealed.
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In autumn, chlorophyll gets degraded and the yellow and orange colours of carotenoids are exposed.
(Credit: wikipediacommons) |
Poisoned
by light
Over the past 20 years,
studies of plants and algae lacking each of the four main xanthophylls showed
they have specific functions in photo-protection. Photosynthesis generates a
waste product without which we couldn’t live: oxygen (O2). But in
the presence of excessive light, toxic forms of oxygen, called reactive oxygen
species (ROS), are produced. Photosynthetic organisms have come up with clever
ways of protecting themselves against photo-damage induced by ROS. Some plants
can simply turn their leaves to minimise light absorption, but most invest in
dissipating excess photons and getting rid of ROS.
Xanthophylls protect the
photosynthetic apparatus from photo-damage by scavenging ROS, by preventing its
formation or by converting the excess light energy into heat. They are mostly
found in the so-called ‘antenna’ protein-pigment structures, which absorb and
transfer light energy to the photosynthetic reaction centres. And this is where
the magic happens: light energy is converted into chemical energy, used to make
organic matter from carbon dioxide removed from the atmosphere.
Without each of the
xanthophylls, plants become photosensitive but are still able to grow. However, a new study led by Bassi and his colleague Luca Dall'Osto shows that simultaneously removing all four xanthophylls,
but not carotenes, has dramatic effects for plants. “If you avoid the synthesis
of xanthophylls the plant is not viable anymore,” Bassi says “It’s something
really new and was completely unexpected.” The researchers blocked the
synthesis of all xanthophylls in Arabidopsis
thaliana, a weed widely used in research, by using a combination of genetic
mutations they called nox (for ‘no xanthophylls’). They found that, in
addition to their role in light absorption and photo-protection, xanthophylls
are essential for plant development- the nox
plants simply couldn’t grow. But this didn’t make sense. Previous studies
showed that removing each xanthophyll at the time, or removing the antenna proteins
to which they bind to, doesn’t affect plant growth. So why do these nox plants have such severe
developmental defects?
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Plants without xanthophylls (right panel) can be 'forced' to grow on a suger-rich medium, but they have severe growth defects and white leaves because they lack photosynthetic complexes. Left: normal plants. (Credit: Luca Dall'Osto) |
A
function at the core
To try and understand this,
the researchers rolled up their sleeves and plunged into some serious
biochemistry. When they analysed the proteins in the two photosynthetic
reaction centres - photosystems I (PSI) and II (PSII) - they faced yet another
surprise: the PSI core proteins had nearly completely vanished in the nox plants. This was strange because
xanthophylls bind mainly PSII antenna proteins, which were mostly unaffected,
but not PSI. “It was so surprising that we spent two years repeating the
experiments to be sure we didn’t make a mistake,” Bassi recalls.
Chloroplasts, the home of photosynthesis, contain about 120
genes encoding for proteins of the photosynthetic machinery. Bassi and
colleagues found that xanthophylls are important for the translation of some of
this genetic information into PSI core proteins, such as the PsaA/PsaB unit,
within chloroplasts. It was known that without the PsaA/PsaB central unit the PSI
super-structure can’t assemble properly and eventually gets degraded, and this is
exactly what the researchers see in the nox
plants.
The absence of a functional PSI
reaction centre in plants lacking xanthophylls explains their developmental
defects, but it remains unclear why this pigment acts specifically on the
synthesis of certain PSI core proteins. It is also difficult to imagine how
xanthophylls, which are embedded in the chloroplast membranes, can act on the
cell protein factories, the ribosomes, dispersed inside the chloroplast. Bassi
believes that xanthophylls are chopped into smaller molecules that leave the
membrane and interact with ribosomes to control the production of PSI core
proteins, and his team is currently searching for these molecules. “There is
some evidence that products from cleavage of other carotenoids are involved in
development, now we know that products of carotenoids are needed for
translation of genes,” he explains.
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Roberto Bassi's research team. (Credit: Luca Dall'Osto) |
An
enlightened career
While part of Bassi’s team
continues investigating carotenoids in Arabidopsis,
the rest studies photosynthesis using his favourite model system: Chlamydomonas reinhardtii. This single-cell green alga is used in photosynthesis
research because it grows faster than plants but their photosynthetic
machineries are nearly identical. After a first postdoc in Copenhagen isolating antenna proteins from barley, Bassi
reckoned that biochemistry would only take him so far in his research, so he
spent the next years learning biophysics with Pierre Joliot in Paris and then molecular
biology with Jean David Rochaix in Geneva, both experts and pioneers in Chlamydomonas photosynthesis research. Bassi
then returned to Italy to set up his own lab to study photosynthesis in algae at
Padua University, but finding little support there for his research he decided
to leave and finally settled at Verona University. The idea of working with
algae was not appreciated by the Italian research funding agencies, however, so
eventually he was forced to switch model system from algae to plants.
Science
funding in Italy: a political tragedy
“The funding situation for
science in Italy is extremely bad and we live with money from Europe,” Bassi
says. His team of 15 people currently runs on three European grants and some
funding from the private sector. For over a decade, under the government of
former Prime Minister Silvio Berlusconi, public research funding and university
recruitments in Italy lacked transparency and a clear strategy. With the
election of a new government in 2011, Italian scientists were hopeful the
situation would improve, but because of the global economical crisis and the
country’s huge public debt, Italy’s modest science budget suffered further cuts
as part of the government’s austerity plan. As a result, despite recent reforms
in the public science funding system, Italian research continues to rely
considerably on European grants. Increasingly aware of this grim situation,
thousands of Italian researchers leave the country every year with little expectations
of going back. Did Bassi ever feel tempted to move abroad? In 2002 he took the
plunge and moved to France to set up a lab in Marseille, but after three years he
had to return to Verona for personal reasons.
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Algae photo-bioreactor (Credit: kaibara87/everystockphoto) |
From algae to biofuels
In
recent years the Italian government began to show some interest in the
production of biofuels from algae, so after a hiatus of many years Bassi could work
with Chlamydomonas again. “I like the field
of bioenergy because the applied research and the basic research are very much
the same thing,” he says. His team studies photosynthesis while trying to
improve the efficiency of biofuel production in algae. Biofuels made from plants, such as corn or soybean, while initially
regarded as a good alternative to fossil fuels like petrol, in reality cause
great damage to the economy. The problem is that these crops take up a lot of
land that would otherwise be used for agriculture, leading to food price
inflation in the long run. Bassi’s team is part of the Sunbiopath international research
consortium funded by the European Union, which aims at optimising biofuel
production by photosynthetic algae for commercialisation. His team has already succeeded
in genetically engineering algae strains that use light energy more efficiently, making them grow faster and thus
generate more biofuel. The researchers are now trying to expand these
results to the industrial scale. “I think this is a promising accomplishment
and I’m very positive we could produce biofuels from algae.”
Reference:
Dall'Osto L., Piques M., Ronzani M., Molesini B., Alboresi A., Cazzaniga S. & Bassi R. (2013). The Arabidopsis nox Mutant Lacking Carotene Hydroxylase Activity Reveals a Critical Role for Xanthophylls in Photosystem I Biogenesis, The Plant Cell, 25 (2) 591-608. DOI: 10.1105/tpc.112.108621
This article was published in Lab Times on 3-05-13. You can read it here.