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future there are many possible avenues that hold out promise for a more sustainable world.
Current Opinion in Biotechnology 2010, 21:271276 This review comes from a themed issue on Energy biotechnology Edited by Jan R van der Meer and Alexander Steinbuchel
World energy use (values for 2005) is approximately 500 exajoules per year (EJ, or 5 1020 J yr1). Of this total, 8090% is derived from fossil fuels [6]. In equivalent units, this is an average consumption of 16 TW (1.6 1013 W). In comparison with 1980, this represented an approximate doubling of energy use over the previous 25 years. On the basis of the earths surface area of $510,000,000 km2 and the value of annual solar radiation (above) one can calculate that the worlds energy needs would be satised with an area of 1,150,925 km2 if 10% of the solar energy could be converted to useable energy [5], i.e. $0.02% of the total surface area or 0.07% of the land surface, i.e. equivalent to a country the size of Ethiopia, Egypt or Colombia. These then are the realistic values that we need to work with in considering solar energy conversion into organic products by cyanobacteria, algae and plants.
Introduction
Oxygenic photosynthesis occurs in cyanobacteria and their eukaryotic successors, algae and land plants and is summarised by the equation CO2 H2 O light energy ! CH2 O O2 (where the parenthesis around CH2O is a reminder that this is only a portion of an organic molecule, in this case a sugar molecule; and the energy required is 480 kJ per mole of CO2)[1,2]. This process is the major route for solar energy conversion on the Earth today and is carried out by cyanobacteria, algae and plants. Other inputs of organic energy into the biosphere (chemical, thermal, etc) are by comparison negligible. So in looking for future replacement of fossil fuels, harnessing solar energy in photosynthesis, either natural or articial, is an attractive option, since there are vast amounts of unused solar radiation. On the negative side, this option has serious problems with efciency, carbon footprint and in its conict with growing food for a hungry world. Nevertheless in the
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Table 1 Budget for solar energy at various sites on the Earths surface and theoretical primary productivity at various latitudes. *Values for the tropic of capricorn and 37-S are similar. The values for daily carbon are for the summer solstice, the equinox and the winter solstice, except for the Equator where there is only one solstice. Taken from [3]. Latitude Equator Darwin (128280 S) Growing season All year (365 d) All year (365 d) Daily carbon xation (gC 2 1) 19.8 17.6 20.3 19.2 13.9 21.8 17.6 9.90 22.3 14.2 4.98 20.8 8.09 0.606 Total Irradiance (E m2) 20 238 19 710 Total Carbon Fixation (gC m2) 6 823 6 602
18 615
6 136
378N*
7 Months (214 d)
12 208
4 031
558N
5 Months (153 d)
8 130
2 533
sion into organic matter by plants and algae yield rates of 18 gC d1 (Table 1), which is an efciency of conversion of $3.2% (cf. [2,9,10]). However, the best solar energy conversion rates of agricultural plants, e.g. Napier grass and sugar cane, are much less than this, with a maximum of 1.6% [2,11,12] and these are the star performers grown under the most pampered conditions: the more usual rates for temperate crops and forests are 0.2 to 0.4%. It is true that only a part of the primary production gives rise to the crop (and only parts of the crop are used). Using all the aboveground parts could improve yields; and thus biomass production of biofuels has potential to improve yields, but probably not much more than, at maximum, a 2.5% conversion rate. This would leave below-ground parts largely unutilised. Cyanobacteria and algae have the further potential that in theory all the primary production could be utilised [10], but other factors mean that algae overall are little more efcient, and in terms of energy input much more costly [13] (also see Photosynthesis in air vs photosynthesis in water and Bioenergy from algal ponds and photobioreactors). Furthermore, in all these possibilities there are many hidden drawbacks, which have hindered realisation of high efciency of conversion, with the attendant burden of high input of energy, largely in the form of fossil fuels, and fertilisers (see, e.g. refs [10,14]). The low yields from plants, algae and cyanobacteria, is a constant frustration in comparison with physical methods of solar energy conversion such as solar photovoltaic cells. It is not likely that genetic engineering will greatly improve the efciency in the short to medium term, although small gains are possible. The major losses of photosynthesis can be put down to some major, as well as many smaller, inefciencies [2,7,8,13]: 1. Two photosystems (in series) rather than one.
Current Opinion in Biotechnology 2010, 21:271276
2. Dependence on a chlorophyll-based system that is dependent (in the greater part) on the spectrum of chlorophyll and on harvesting energy from the rstexcited singlet state of chlorophyll a. 3. Inefciencies of light-harvesting, levels of enzymes vs. changing light climates, and chemical inefciencies. 4. Photoinhibitory damage to photosystem II. To re-engineer natural photosynthesis to overcome these inefciencies will not be an easy challenge and has led to a move to biomimetics of photosynthesis, i.e. towards articial photosynthetic systems (see Hydrogen production and articial photosynthesis).
Photosynthesis in air vs photosynthesis in water: Land vs ponds and lakes vs the sea
Land plants have a hidden advantage over aquatic plants in that they have evolved homoiohydry to enable them to photosynthesise in air using leaves that have evolved to utilise the rapid diffusion of gases in air, which is of the order of 10,000 times faster than that in water [15,16,17,18]. Thus algae or cyanobacteria (in water) must either be limited by stirring and grow slowly - giving rise to a low rate of natural primary production - or be stirred and grow faster, which for algal ponds and photobioreactors means employing some form of mechanical stirring (see Bioenergy from algal ponds and photobiorectors). Primary production by phytoplankton in lakes or oceans is very low [9,11]. The major reasons for this are because (i) stirring in such waters is low, and (ii) levels of essential nutrients are also low. Nevertheless, the large areas involved and the essentially natural and free primary production might make this an attractive area for xedcarbon harvesting; however, the difculty of harvesting the product makes this is an unacceptable option, except for shallow ponds.
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Limitations and prospects of natural photosynthesis for bioenergy production Larkum 273
In shallow lagoons and ponds, especially those that are polluted, opportunities do exist [19]. Nevertheless the difculties of adapting new practices to shallow water bodies will certainly delay this use for a number of years. As a result, terrestrial plant bioenergy will be most used in the near future (see next section).
deserves. Other problems are photoinhibition and other stresses [56] decline in yields due to pathogens, ageing, etc, and optimising optical path lengths and the package effect, as well as the large scale nature of such projects if they are to meet the needs of bioenergy production [10]. Thus, while these systems present novel advantages over land-based agriculture and water-based mariculture, their potential must be thoroughly assessed and the disadvantages acknowledged. Theoretical yields are discussed in Solar energy conversion by cyanobacteria, algae and plants. There will be much debate for some time to come over the actual yield potentials from algae. This is because in single celled algae it should be possible to redeploy a greater fraction of absorbed solar energy into xed carbon compared with land plants with their stems and root systems. However, the culturing of algae brings with it another set of problems such as stirring, nutrient supply, optimisation of the light eld, optimisation of the growth conditions and protection from pathogens and nuisance weed algae. While these factors have been discussed for small-scale pilot plants by many workers in recent times [5662] there is no clear way forward. Another area of confusion is over what is a realistic cropping factor; and on this there is at present a wide range of opinion for algal pond and photobioreactors; but little good data. Small-scale facilities can be nurtured to yield impressive xed-carbon outputs. However, it remains to be proved that this can be done on a long-term basis at commercial levels of at least 1000 ha. Even then it must be shown that transport costs to the area can be economically viable. Finally, water in many regions may be a prime commodity and the requirement for large amounts of water for algal ponds or photobioreactors may be antithetical to the overall need to conserve water. In the short-term, the prospect for the use of algae to provide signicant amounts of organic carbon, to sustain the human need for bioenergy, is limited. In the longer term the possibilities should be explored, but they come at the cost of a high solar footprint (see below), a high carbon footprint and many unknowns [63]. Over the recent past there have been some notable failures in this challenging eld [64]. Initially it can be seen that small-scale success can be obtained by taking advantage of local circumstances. For example, heat, stirring and nutrients could be obtained from power stations. Coastal lagoons and other shallow water bodies could be advantageous for algal ponds. Initially these could be operated with little stirring and no nutrient input, and harvesting could be done by simple means. In the longer term solar photovoltaic or solar thermal assemblies could be used to provide much of the energy needed for the algal farms. Sewage and other efuents could be used to provide nutrients.
Current Opinion in Biotechnology 2010, 21:271276
Perhaps the brightest possibility as regards algae is the production of value-added products. The commercial production of beta-carotene [65] is a good example here. There are many other potential products from algae such as sugars, biopolymers, lipids and antibiotics. In negotiating the commercial output of these products, the algal biotechnology industry would learn much that could eventually be fed back into the system in the production of algal biofuels.
more if solar photovoltaic panels are contemplated to offset the high-energy needs of algal farms, the solar and carbon footprints greatly increase.
Conclusions
Natural photosynthesis is an ancient process on the Earth that evolved under conditions where efciency was not a selection pressure. The most efcient photosynthetic organisms are land plants. Algae and cyanobacteria, living under light-limited conditions, are generally less efcient but offer possibilities in the future. The 12% efciency of plants, algae and cyanobacteria cannot compete with solar photovoltaic panels or solar thermal collectors. Nevertheless natural photosynthesis will become increasingly used for the production of biofuels and organic products in a world where these will increasingly come into short supply. It seems unlikely that algae and cyanobacteria will be used as a source of bioenergy. In the longer term photobiohydrogen production and articial photosynthesis hold out possibilities. In all these cases, high carbon footprints, land sequestration, loss of biodiversity and other problems must be addressed.
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