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Turkish Journal of Biology Turk J Biol (2015) 39: 276-283 © TÜBİTAK doi:10.3906/biy-1406-62 http://journals.tubitak.gov.tr/biology/ Research Article Energy efficiency of the sunlight harvesting and storing system in bacterial photosynthesis: comparison with semiconductor photovoltaic cells 1, 2 1 1 Erkin ZAKHIDOV *, Mavluda ZAKHIDOVA , Abdumutallib KOKHKHAROV , Abdurasul YARBEKOV , 1 1 1 1 Vakhobjon KUVONDIKOV , Sherzod NEMATOV , Erkin NORMATOV , Aziz SAPARBAYEV 1 Institute of Ion Plasma and Laser Technologies, Academy of Sciences of Uzbekistan, Tashkent, Uzbekistan 2 National University of Uzbekistan, Tashkent, Uzbekistan Received: 21.06.2014 Accepted: 15.10.2014 Published Online: 01.04.2015 Printed: 30.04.2015 Abstract: The light phase of photosynthesis is considered a joint operation of 2 functional pigment–protein complexes: a light harvesting antenna, absorbing sunlight in a wide spectral range, and a reaction center, utilizing the energy of absorbed light quanta in photochemical charge separation reactions. These complexes allow converting solar energy to the energy of specific biomolecules with high quantum efficiency. However, before being transferred to reaction centers, the solar energy is stored in the lowest excited state of pigment molecules of the light harvesting antenna that partially convert light quantum energy into heat. These energy losses bring a significant reduction of energy efficiency of photosynthesis in view of a very wide spectral range of photosynthetically active sunlight. In the current study we analyzed the energy efficiency of sunlight harvesting and storing in different photosynthetic bacteria with different absorption bands. We showed that simultaneous exploitation of several such photosynthetic organisms leads to an increased total energy efficiency in terms of harvesting sunlight of the wider spectra. Maximal values of energy efficiencies of the sunlight harvesting and storing system in photosynthetic bacteria and semiconductor photovoltaic cells are compared, and perspectives on practical use of photosynthetic bacteria as solar energy converters are discussed. Keywords: Sunlight, plant and bacterial photosynthesis, light harvesting and storing system 1. Introduction Sunlight is the main energy source on earth. The total energy of sunlight striking the earth’s surface at a rate of 1017 W is more than sufficient to meet all of mankind’s energy needs, currently estimated at 1.5 × 1013 W (US Department of Energy, 2013). However, there are 2 main problems limiting the effective utilization of sunlight: it is a rather diluted source of energy, and it has a very wide spectral range. At 90° latitude on the upper border of the earth’s atmosphere, sunlight has an intensity of 1350 W/ m2, and at sea level its intensity is approximately 1000 W/m2. Taking into account dark periods and alterations of sunlight flow at different latitudes, the annual average sunlight intensity on the earth’s surface is less than 170 W/m2 (Lewis and Nocera, 2006). In Figure 1, the sunlight spectra measured on the upper border of the earth’s atmosphere and at sea level (Barber, 2007) are shown. Comparing these 2 spectra, one can notice that earth’s atmosphere significantly reduces sunlight intensity in the short-wavelength range. In the long-wavelength range there are specific narrow bands due to strong absorbance *Correspondence: [email protected] 276 of sunlight (absorption peaks of O2 and H2O) (Gueymard, 2004). Thus, sunlight energy is basically concentrated in the spectral range of 350–1200 nm. Excluding low-potential thermal utilization of sunlight energy, the most advanced device for sunlight energy conversion with highest efficiency is the semiconductor photovoltaic (PV) cell. Recently a 44.7% conversion efficiency in a 4-junction PV structure was announced (Fraunhofer Institute for Solar Energy Systems, 2013). However, there are 2 essential factors potentially limiting overall application of such structures: 1) the electrical current generated by a PV converter changes during the day (and the year) depending on the intensity of striking sunlight and stable delivery of such energy to consumers requires proper accumulation, which is costly and technically problematic; and 2) the production of PV electrical power comparable with that of traditional electric stations requires an increase in manufacturing of semiconductor materials, which poses significant environmental risks because of the specificity of chemical materials and technologies used in the process (Gottesfeld and Cherry, 2011). ZAKHIDOV et al. / Turk J Biol 10 Sunlightintensity, a.u. On the upper border of the atmosphere On the sea level 5 0 350 450 550 650 750 850 Wavelength, nm 950 1050 1150 Figure 1. Sunlight spectra measured on the upper border of the earth’s atmosphere and at sea level. The characteristic absorption peaks are caused by O2 and H2O and are located in the NIR region (Barber, 2007). A natural alternative to the PV conversion of solar energy on the global scale might be photosynthesis (PS), a complex of photochemical reactions in plant leaves or PS bacteria driven by sunlight. In PS systems fast photochemical processes transform sunlight energy into chemical energy of specific biomolecules (ATP and NADP+) through the energy of a transmembrane hydrogen gradient (electric potential). These processes exhibit a high rate of quantum efficiency (up to 100%) (Ruban, 2013). Moreover, during the slow dark reactions of the Calvin cycle this energy is stored in a much more stable form as glucose or its derivatives (Clayton, 1984). Being controlled by specific enzymes, these reactions depend on the physiological state of PS organisms and may be essentially inhibited by environmental stressors such as drought and salt stress, sub- or supraoptimal temperatures, and excessive illumination (Zakhidov et al., 2002; Fracheboud and Leipner, 2003; Culha Erdal and Cakirlar, 2014). It should be noted that PS organisms have effective mechanisms for self-protection from such stressors. For example, plants exposed to excessive sunlight may change the structure and content of PS pigments to prevent the destruction of reaction centers through dissipation of excessive energy of absorbed light into heat (Zakhidov et al., 2006; Murata et al., 2007) or they may limit dehydration by leaf rolling (Terzi et al., 2013). Unlike PV cells, PS organisms have another important function: replication, which utilizes additional resources and reduces the total energy efficiency of PS (Pace, 2001). If PS energy efficiency is measured as a ratio of the burning energy of the biomass accumulated by plants during the vegetation period (or a 1-year period) to the sunlight energy absorbed by them for the same period, it does not exceed 1% even in crop plants grown under tropical conditions (Barber, 2009). High efficiencies can only be reached in algae grown in special bioreactors (Janssen et al., 2003; Wijffels and Barbosa, 2010). From the total sunlight spectral range of 350–1200 nm (Figure 1), the part with wavelengths of less than 700 nm is utilized in both plant and bacterial PS, whereas the near infrared (NIR) part with wavelengths of 700–1100 nm is used only in bacterial PS (Sheer, 2012). Plant PS is often considered as a practical and easy-to-use system for utilization of solar energy because of efficient biomass production. The advantages of bacterial PS are related to both operations in the NIR spectral range having up to 40% of total sunlight energy and the possibility of genetic tailoring of PS bacteria for optimized light conversion (Bylina et al., 2002). Furthermore, unlike plant PS with its 2 photosystems and twice reduced energy efficiency of sunlight conversion due to nearly the same absorption spectra (Blankership et al., 2011), bacterial PS with a single photosystem has no such energy efficiency reductions. Spatially and functionally divided but effectively cofunctioning, the light harvesting antenna (LHA) and the reaction center (RC) are 2 pigment–protein complexes that provide the highest quantum efficiency of photochemical processes. However, the energy of sunlight quanta absorbed in the LHA is stored in the lowest excited state of PS pigment molecules before being transferred to the RC, thereby partly losing the sunlight quantum energy to heat (Ruban, 2013). As observed in PV cells, these losses cause a corresponding decrease in energy efficiency of PS conversion. The lower energy of the excited state causes decreased energy efficiency of the PS conversion. Various genotypes of PS bacteria even from the same family may have extremely different absorption spectra. Their long-wavelength bands undergo essential spectral shifts depending on the structure and molecular environment of the PS pigments. In the case of simultaneous exploitation of a number of such PS bacterial cells, their common absorption spectra may be considerably extended. Such multicellular, aggregated bacterial PS systems may be considered as analogues to multijunction PV cells (Green et al., 2011). In the present paper, spectral characteristics of a group of PS bacteria with different absorption bands in the NIR were studied. Absolute values of energy efficiencies of sunlight harvesting and storing in these PS bacteria were calculated using the proposed simplified model of 2-band absorption spectra in separate and joint operations of PS bacteria. We propose a novel system with increased energy efficiency of sunlight harvesting and storing composed of multicellular aggregated PS bacteria with different spectral forms and modified absorption spectra. Energy efficiencies of sunlight conversion in PV and in PS are comparatively analyzed. 277 ZAKHIDOV et al. / Turk J Biol 3. Results In Figure 2, the absorption spectra of solutions of whole cells (Figure 2a) and extracted PS membranes (Figure 2b) of B. viridis at room temperature are shown. The comparison of these 2 spectra clearly shows that in whole cells with sizes of approximately 1 µm, light scattering losses are very high, especially in the short-wavelength range. In the solution of extracted PS membranes with much smaller sizes, light scattering losses are much lower, allowing measurement of the spectral characteristics of absorption by PS pigments with higher accuracy. Further dissociation of the PS system induces both essential spectral shifts of relevant absorption bands and changes in their function (Parkes-Loach et al., 1994). In Figure 3, room temperature absorption spectra of solutions of PS membranes extracted from 5 wild-type PS bacteria are shown. At low concentrations of PS membranes, 278 1.5 Absorption. a.u. 2. Materials and methods Absorption spectra of wild-type purple PS bacteria (Blastochloris viridis, Rhodospirillum rubrum, Rhodobacter sphaeroides), green-sulfur bacteria (Chloriobium phaeobacteriodes), and cyanobacteria (Synechocystis) were studied (PS membranes extracted from wild-type purple PS bacteria were kindly provided by Prof JR Norris and Dr NS Ponomarenko from the University of Chicago, Chicago, IL, USA). The cells of the studied bacteria were cultivated at 20–30 °C and under illumination of 10 to 25 W/m2 using methods described by Madigan and Jung (2008). Light intensity in all experiments was measured by Lux-Meter Yu-116, sensitive in the visible and NIR spectral ranges. For the exact establishment of peak wavelengths and amplitudes of relevant absorption bands, measurements were carried out with extracted PS membranes of bacteria in low concentration solutions, when the extinction coefficient of the main IR band was less than 1.0. Photochemically active membranes were extracted from whole cells of bacteria using low-frequency sonication and subsequent ultracentrifugation (Ponomarenko et al., 2010). This allowed preserving the nativeness of PS membranes (despite the essential structural disintegration of cells) and the proceeding relevant photochemical reactions (Beatty et al., 2005). Light scattering losses in solutions of PS membranes were much lower than those in solutions of whole cells, which allowed a more accurate measurement of the net absorption spectra. The absorption spectra of PS membranes in physiological solution of pH 6.9 at a room temperature of about 22 °C were measured in the spectral range of 350– 1100 nm using a Shimadzu 1601 spectrometer (Shimadzu Europa, GmbH, Duisburg, Germany). The intensity of the measuring light in all experiments was less than 10 W/m2, a level that does not affect the absorption amplitudes and spectra. 1 a 0.5 b 0 350 450 550 650 750 850 950 1050 Wavelength. nm Figure 2. Room temperature absorption spectra of low concentration (extinction coefficients of the main IR bands ≤1.0) solutions of whole bacteria (a) and extracted PS membranes (b) of B. viridis. It is clearly seen that extraction of PS membranes improves the exact definition of wavelengths and amplitudes of absorption bands in the long-wavelength region. when the QX absorption band of bacteriochlorophyll (BChl) molecules associated with PS membrane, as well as all absorption bands of monomer BChl molecules (Olson and Stanton, 1991), may be neglected, each absorption spectrum has 2 main bands. The short-wavelength band in the range of 350–550 nm is composed of a Soret band of BChl and a 3-peak-band of carotenoids. The longwavelength band is the QY absorption band of BChl with a spectral width of approximately 100 nm and varying in the range of 720–1020 nm depending on the type of BChl molecule (a, b, or d) and the molecular surrounding of the PS membrane (Scheer, 2006). Thus, as a simplified model of absorption spectra of PS membranes, it is possible to accept these spectra composed of 2 bands: a wider shortwavelength band in the range of 350–550 nm and a narrower, approximately 100 nm, long-wavelength band with a varying spectral position in the range of 720–1020 nm depending on the genotype of the PS bacteria. In B. viridis (curve 1) with BChl b as the main PS pigment, the QY absorption band (80 nm in width with a peak wavelength of λQY = 1020 nm) had the highest frequency shift of 2840 cm−1 relative to the QY absorption band of monomer BChl b (λQY = 791 nm). Such a gigantic shift of BChl b in the PS membrane is related to the specific features of interaction of this molecule with α- and β-polypeptides in the LHA protein of B. viridis and the structure of the BChl b molecule in the form of tetrapyrrole rings with a long phytol tail (Grimm, 2006). It should be noted that the spectral position of the QY absorption band is sensitive to the environment. Changes in ambient temperature, pressure, or content of surrounding proteins may cause changes in the spectral position of this band Absorption, a.u. ZAKHIDOV et al. / Turk J Biol 5 4 1 0 350 450 550 1 3 2 650 750 850 Wavelength, nm 950 1050 1150 Figure 3. Room temperature absorption spectra of PS membranes of wild-type B. viridis (1), R. rubrum (2), R. sphaeroides (3), C. phaeobacteriodes (4), and cyanobacterium Synechocystis (5). Each spectrum is normalized by its main IR band. Joint functioning of all these PS samples allows harvesting and storing of almost the entire spectrum of active photosynthetic sunlight radiation. (Freiberg et al., 1993; Monshouwer et al., 1995; Zakhidov et al., 2004). In R. rubrum (curve 2) the main PS pigment, BChl a, has a QY absorption band (60 nm in width with a peak wavelength of λQY = 873 nm) with a shift of 1500 cm−1 relative to that in monomer BChl a (λQY = 772 nm) (Olson and Stanton, 1991). Although this frequency shift is smaller than that of B. viridis, it is still high and relates to a strong interaction of the BChl a molecule with surrounding proteins in the PS membrane. R. sphaeroides, unlike the 2 above-mentioned PS organisms, has both core (λQY = 875 nm and peripheral (λQY = 800 nm and 850 nm; curve 3) antennas. The main PS pigment (BChl a) in these bacteria has 3 spectral forms with a total width of 115 nm of the QY absorption band: 2 expressed peaks with frequency shifts of 1525 cm−1 and 1190 cm−1, and a shoulder peak in the longer wavelength part of the spectrum with a frequency shift of 450 cm−1 relative to the spectral position of monomer BChl a. The presence of a peripheral antenna having a BChl a QY absorption band with higher energy allows R. sphaeroides to harvest additional sunlight and transfer sunlight energy to BChl a of the core antenna (Abdurakhmanov et al., 2013). C. phaeobacteriodes, a green-sulfur PS bacterium, has BChl d as the main pigment with λQY = 654 nm in monomer form. In the PS membranes of this bacterium the QY absorption band (45 nm in width and λQY = 720 nm, curve 4) is shifted 1400 cm−1 relative to that of monomer BChl d (Montesinos et al., 1983). Depending on light intensity and spectrum, the structure of pigments and the spectral position of their absorption bands in C. phaeobacteriodes may undergo dramatic changes, called biofiltration of light (Parkin and Brock, 1980). Cyanobacterium Synechocystis has 2 key differences from the above-described PS bacteria: the main PS pigment is chlorophyll a (Chl a), but not BChl, and there are 2 simultaneously operating photosystems, as in higher plants, providing molecular oxygen (Barry et al., 1994). The energy of sunlight absorbed in the spectral range of 350–500 nm is, basically, transferred to Chl a through phycobilisomes. There are 2 spectral forms with maxima at 630 nm and 670 nm, and a total spectral width of approximately 140 nm (curve 5). These chlorophylls accordingly further activate photochemical reactions in the 2 photosystems (Shestakov and Mikheeva, 2006). In wavelengths longer than the wavelengths of the appropriate QY absorption bands, all of the abovedescribed PS bacteria are transparent; thus, they will not shade each other and all together may harvest sunlight in much wider sunlight spectra. In Figure 4, comparative energy diagrams of sunlight harvesting and storing by PS bacteria B. viridis (left side) and crystalline silica PV cells (right side) are shown. In the PS membranes of B. viridis sunlight quanta with wavelengths of 350–1050 nm (or energies of 1.1 ÷ 3.5 eV) are absorbed in a very short time (~10−12 s) and stored in the lowest excited state of BChl b with an energy of 1.22 eV (or a wavelength of 1020 nm). There is a partial loss of energy of light quanta that will further drive the photochemical reaction in the RC. Generally, the same process takes place in a crystalline silica PV cell: light quanta with energy higher than the energy of the forbidden band (or with wavelengths shorter than the threshold wavelength) transfer electrons from the valence band to the conductance band. Thus, the energies of these electrons have to be different, depending on the energies of the light quanta driving such an internal photoeffect. These electrons rapidly relax to the bottom of the conductance band to initiate a photoelectric effect (Dorofeev et al., 2009). Despite the high quantum efficiencies of these 2 processes, a significant part of the energy of sunlight quanta gets lost in the transferring of light excitation to the lowest excited state of BChl b in B. viridis. An energy reduction of sunlight quanta also happens in crystalline silica by the relaxation of electrons to the bottom of the conductivity band, which causes rather low energy efficiency of light harvesting and storing in the PV cell. High values of energy efficiency in semiconductor PV cells may be achieved in multilayer semiconductors with different forbidden bands (Nelson, 2003; Bosi and Pelosi, 2007). The same approach may be used for creating high efficiency sunlight converters based on various PS systems with different absorption bands. 279 ZAKHIDOV et al. / Turk J Biol E , (eV ) E , (eV ) E , (eV ) Soret band Absorption bands of additional pigments Conductance band Q х b and Q у band The basic state of BChl Valence band Figure 4. Comparative energy diagrams of sunlight conversion in B. viridis (а) and silica PV cells (b). Bold straight lines with arrows mean electronic transitions driven by light quanta; waved lines mean heat relaxation from high excited states to the lowest excited state with a partial loss of quantum energy; and dotted lines mean transitions converting sunlight energy to energy of PS or PV, accordingly. Absorption spectra of these 2 samples are shown on the left side of each diagram and the spectrum of sunlight striking the earth’s surface is in the middle. Energies of light quanta are presented in eV. 4. Discussion Energy efficiency of PS light harvesting and storing may be qualitatively estimated using a simplified model of uniform spectral distribution of sunlight intensity. In this model, sunlight energy in a certain spectral range is proposed to be proportional to its spectral width. In the short-wavelength range, B. viridis PS membranes absorb mostly at wavelengths of 350–550 nm. Taking into account that the sunlight spectrum extends in the range of 350– 1200 nm, the proportion of sunlight absorbed by B. viridis PS membranes will be (550 nm – 350 nm)/(1200 nm – 350 nm) = 0.24. The energy efficiency of sunlight storing in this case is determined as a ratio of energies of stored quanta (in the lowest excited state of BChl b with an energy of 1.22 eV) and absorbed quanta (in the short-wavelength band of 350–550 nm, with an average energy of 2.9 eV), which is 0.42. In the long-wavelength band, the QY band, with a width of 80 nm, sunlight with a proportion of 80 nm/ (1200 nm – 350 nm) = 0.095 will be stored with an energy efficiency of 1.0, since there is no sunlight quanta energy 280 loss in this case. In total, light harvesting and storing in B. viridis has the following energy efficiency: (0.24 × 0.42) + (0.095 × 1.0) ≈ 0.20. In R. rubrum the proportion of sunlight absorbed in the short-wavelength range of 350–550 nm is the same: (550 nm – 350 nm)/(1200 nm – 350 nm) = 0.24. The sunlight absorbed with an average wavelength of 873 nm will be stored on the BChl a QY band at 873 nm (energy losses are ~ 2 times and, thus, energy efficiency is ~ 0.5). Furthermore, sunlight quanta harvested and stored in the same BChl a QY absorption band has a proportion of 60 nm/(1200 – 350 nm) = 0.07 and an energy efficiency of 1.0. Therefore, the total energy efficiency in R. rubrum is (0.24 × 0.51) + (0.07 × 1.0) ≈ 0.19. R. sphaeroides has the same spectral position of the short-wavelength absorption band as R. rubrum, but a wider spectrum of the 3-peak QY band. In this case the energy efficiency of harvesting and storing shortwavelength sunlight is 0.51. However, the wider QY band of R. sphaeroides provides a higher proportion of sunlight ZAKHIDOV et al. / Turk J Biol harvesting in the long-wavelength range: 115 nm/(1200 nm – 350 nm) = 0.14. As a result, the total energy efficiency of sunlight harvesting and storing in R. sphaeroides is (0.24 × 0.51) + (0.14 × 1.0) ≈ 0.26. In C. phaeobacteriodes, sunlight is absorbed in the range of 350–550 nm and stored in the QY band with a shorter wavelength, λQY = 720 nm, with an energy of 1.72 eV. Thus, these sunlight quanta lost less energy and the energy efficiency of their storing is 0.59. The QY absorption band in C. phaeobacteriodes is narrower than those in other PS bacteria (45 nm), and that is why the proportion of the absorbed long-wavelength sunlight is less: 45 nm/ (1200 nm – 350 nm) = 0.05. In total, the energy efficiency of sunlight harvesting and storing is (0.24 × 0.59) + (0.05 × 1.0) ≈ 0.19. The proportion of sunlight absorbed by cyanobacterium Synechocystis in the short-wavelength range is (500 nm – 350 nm)/(1200 nm −350 nm) = 0.18. It is stored on the QY absorption band at λQY = 650 nm with a partial loss of quantum energy from 3.01 eV (on average) to 1.91 eV. Thus, the energy efficiency in this case is 0.64. In the long-wavelength range the sunlight absorbed and stored in the QY absorption band has the proportion of 140 nm/ (1200 nm – 350 nm) = 0.17. The total energy efficiency of sunlight harvesting and storing in Synechocystis, taking into account the presence of 2 photosystems, is 0.5 [(0.18 × 0.64) + (0.17 × 1.0)] ≈ 0.14. If solutions of the described cells of PS bacteria were to be arranged in sequential positions, with each following PS-type bacterium having longer wavelengths of the QY absorption band than the previous one, mutual shading of these PS bacteria would be minimized and a common energy efficiency of sunlight harvesting and storing in such a multi-PS bacterial system may be easily estimated. In the short-wavelength range the common sunlight harvesting and storing efficiency of the system is roughly the same as in the first PS bacteria with the lowest λQY (cyanobacterium Synechocystis), plus the efficiency of the C. phaeobacteriodes absorption band at 500–550 nm with a proportion of (550 nm – 500 nm)/(550 nm – 350 nm) = 0.25. In the long-wavelength range all described bacteria will additively contribute efficiencies related by their QY bands. Thus, in total, the value of the common energy efficiency of sunlight harvesting and storing in the proposed composed system can be calculated as 0.51. PS bacteria are characterized with extremely high (up to 100%) quantum efficiency of the primary light phase of photosynthesis. However, harvesting of sunlight in a very wide spectral range of 350–1100 nm significantly decreases the PS energy efficiency in these organisms due to subsequent storing in the lowest excited state of the PS pigments. In the different types of PS bacteria studied here, the energy efficiency of sunlight harvesting and storing has been estimated at ~0.20. In order to significantly increase the sunlight absorption spectral range and the efficiency of its simultaneous storing, a multi-PS bacterial system composed of a group of PS bacteria strains with different absorption spectra is proposed. This will increase the energy harvesting and storing efficiency to higher than 0.5. Further enhancement of the PS energy efficiency is possible by optimizing the absorption spectra of PS bacteria. The simultaneous operation of various PS bacterial strains is possible if their physiological and optical behaviors are compatible (Overmann and Garcia-Pichel, 2006). In such conditions a multi-PS bacterial system may be used as an effective convertor of solar energy that allows working out the energy of the transmembrane potential and of the specific carriers of bioenergy such as ATP and NADP+. It should also be noted that the biological origin of the proposed system provides significant advantages from an ecological point of view (Atsumi et al., 2009; Blankership et al., 2011). In conclusion, in the current study, a group of PS bacteria absorbing sunlight in different spectral ranges was studied. It was demonstrated that the simultaneous employment of several PS bacteria results in a considerable increase of the total energy efficiency of such a system by widening the sunlight spectrum utilized in photosynthesis. The comparative study of energy efficiency of sunlight harvesting and storing in PV and PS systems has revealed a number of their common and distinguishing properties. Advantages of PS bacteria for sunlight energy conversion in the NIR region were shown. Acknowledgments This work was completed in the framework of project # F2 – FА – F147 of the Program of Basic Research of the Academy of Sciences of Uzbekistan. The authors acknowledge Prof IY Abdurakhmonov for a critical discussion of the paper. References Abdurakhmanov U, Zakhidov EA, Zakhidova MA, Quvondikov VO, Nematov SQ, Ponomarenko NS, Saparbaev AA (2013). Migration of light excitation in LHA of photosynthetic bacteria: the molecular structure providing high efficiency of energy conversion. 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