Ambio 2016, 45(Suppl. 1):S24–S31 DOI 10.1007/s13280-015-0730-0 From first generation biofuels to advanced solar biofuels Eva-Mari Aro Abstract Roadmaps towards sustainable bioeconomy, photosynthesis. In fact, the line between renewable biofu- including the production of biofuels, in many EU els and non-renewable fossil fuels is sometimes vague and countries mostly rely on biomass use. However, although only complete life-cycle analyses in the future will reveal biomass is renewable, the efficiency of biomass production which feedstocks are truly renewable to be used in biofuel is too low to be able to fully replace the fossil fuels. The production. use of land for fuel production also introduces ethical Depending on the origin and production technology of problems in increasing the food price. Harvesting solar biofuels, they are generally called as the first, second and energy by the photosynthetic machinery of plants and third generation biofuels (according to the EASAC report autotrophic microorganisms is the basis for all biomass 2012), while the fourth generation biofuels (as assigned in production. This paper describes current challenges and the present paper) make use of novel synthetic biology possibilities to sustainably increase the biomass production tools and are just emerging at the basic research level. On and highlights future technologies to further enhance the way towards sustainable bioeconomy, the development biofuel production directly from sunlight. The biggest of efficient biofuel production strategies based on solar scientific breakthroughs are expected to rely on a new energy is of pivotal importance. This concerns EU in technology called ‘‘synthetic biology’’, which makes particular, due to limited bioresources. Despite huge engineering of biological systems possible. It will enable advances in photovoltaic technologies for solar energy direct conversion of solar energy to a fuel from conversion to electricity, the biofuels are still of primary inexhaustible raw materials: sun light, water and CO2.In importance in modern societies, particularly in the trans- the future, such solar biofuels are expected to be produced port sector. In this regard, biofuels have become the largest in engineered photosynthetic microorganisms or in renewable fuel produced and consumed in the world due to completely synthetic living factories. increasing demands to replace the fossil fuels with renewable ones, thus reducing greenhouse gas emissions Keywords Advanced biofuel Á Biomass Á and alleviating climate change. On the other hand, there are Photosynthesis Á Photosynthetic microorganism Á serious concerns about competition of land for food and Solarfuel Á Synthetic biology fuel production, particularly due to the fact that the current technology for biofuel production depends on feedstock derived from the edible fraction of food plants (corn, INTRODUCTION rapeseed, sugar beet and others). Scientific breakthroughs in biofuel production are of key importance in paving the Biofuels are derived from biological material, presently way towards carbon–neutral economy and sustainable mainly from plants, microorganisms, animals and wastes. biofuel production, including both the quantity and quality All biofuels have the same basic and renewable origin. of biofuels for different transport sectors. They are based by the ‘‘present-day’’ photosynthetic con- Today, most of the first generation biofuels are sourced version of solar energy to chemical energy, which sets from crop plants as energy-containing molecules like them apart from fossil fuels that are based on ancient sugars, oils and cellulose. They provide only limited Ó The Author(s) 2016. This article is published with open access at Springerlink.com 123 www.kva.se/en Ambio 2016, 45(Suppl. 1):S24–S31 S25 biofuel yields and have a negative impact on food security. commercial scale to meet advanced non-cellulosic renew- Efforts are now needed to accelerate the generation of able fuel targets, there is a lot of criticism concerning the advanced biofuels by identifying and engineering effective environmental impact, among others. The EASAC report non-food feedstocks, improving the performance of con- (2012) urges systematic research into the anticipated version technologies and the quality of biofuels for dif- improvements from second generation biofuels sourced ferent transport sectors as well as bringing down the costs from inedible parts of plants and recommends further (EASAC 2012). development of third generation biofuels extracted from The second generation biofuels are already an algae as an appropriate means to enhance biofuel produc- improvement in producing biofuels from feedstock of lig- tion. However, far more innovative solutions—the fourth nocellulosic, non-food materials that include straw, generation biofuels or direct solar biofuels and synthetic bagasse, forest residues and purpose grown energy crops on biology technologies—are pertinently needed for replace- marginal lands. Projects are needed to maximise the ment of all fossil fuels. Bioengineering of photosynthetic amount of renewable carbon and hydrogen that can be microorganisms towards direct solar fuel production, i.e. converted to fuels from ‘‘second generation’’ biomass. the fuel production without a biomass phase, has a history The third generation biofuels are based on algal biomass of over 10 years. However, only recently the tools of production. They are presently under extensive research in synthetic biology for efficient and thorough engineering of order to improve both the metabolic production of fuels novel metabolic pathways for fuel and chemical produc- and the separation processes in bio-oil production to tion, as well as optimising the host, have become available remove non-fuel components and to further lower the also for photosynthetic microorganisms. production costs. The fourth generation biofuels—photobiological solar fuels and electrofuels—are expected to bring fundamental EXPECTED SCIENTIFIC BREAKTHROUGHS breakthroughs in the field of biofuels. Technology for IN PRODUCTION OF PHOTOSYNTHESIS-BASED production of such solar biofuels is an emerging field and BIOFUELS based on direct conversion of solar energy into fuel using raw materials that are inexhaustible, cheap and widely Improvement of photon-to-fuel conversion efficiency available. This is expected to occur via revolutionary (PFCE) development of synthetic biology as an enabling technol- ogy for such a change. The synthetic biology field is still in Term PFCE is used here to describe the percentage of the its infancy and only a few truly synthetic examples have energy of photons hitting the organism, converted to been published thus far (see Cameron et al. 2014 for a chemical energy by photosynthetic light reactions and review). For successful progress, one needs to discover subsequently, via a number of processes, ending up in the new-to-nature solutions and construct synthetic living fuel. This terminology was chosen to make some rough factories and designer microorganisms for efficient and comparisons between the efficiency of production of direct conversion of solar energy to fuel. Likewise, a electricity by solar cells (Ingana¨s and Sundstro¨m 2016) and combination of photovoltaics or inorganic water-splitting the production efficiency of biofuels by a number of dif- catalysts with metabolically engineered microbial fuel ferent production systems. production pathways (electrobiofuels) is a powerful The photon-to-fuel conversion efficiency—including the emerging technology for efficient production and storage costs of biomass processing—by the most ‘‘sustainable’’ of liquid fuels. current feedstocks stands only at 0.16 % for Brazil sugar- cane ethanol and ca. 0.15 % for biodiesel from palm oil (see Gouveia 2011) These percentages are much higher CURRENT SITUATION than those for transport biofuels from European wheat and rapeseed. Current research aims at higher PFCEs and better First generation biofuels—direct food-crop-based biofu- quality as absolute requirements for sustainable biofuel els—are likely to be largely banned in EU. In its rigorous production. Developments of various second and third study, EASAC finds that ‘first generation’ biofuels, once all generation biofuels, for example, with improvements in impacts of biomass cultivation are taken into account, biomass processing technologies and by search for better appear to provide little or none of the greenhouse gas feedstocks, like algal biomass with inherently higher pho- reductions required in the Directive whilst putting food, ton-to-biomass conversion efficiency than in land plants, agriculture and natural ecosystems at risk (EASAC 2012). are important steps in this direction. Fourth generation Although the Brazilian sugarcane ethanol is regarded by biofuels are likely to bring much higher improvements in some specialists as the only biofuel currently produced at the PFCE. For example, the future photobiological solar Ó The Author(s) 2016. This article is published with open access at Springerlink.com www.kva.se/en 123 S26 Ambio 2016, 45(Suppl. 1):S24–S31 fuel production system aims at using photosynthetic yield and quality of plant products. Optimising the overall microorganism only as a ‘‘catalyst’’: it harvests the solar energy efficiency of plants/algae and focusing on produc- energy and uses this to direct production of a high-quality tion of the ‘‘most suitable target molecules in planta’’ for fuel in high yield. In optimal cases, the microorganism will biofuel production