Balancing the energy flow from captured light to biomass under fluctuating light conditions

The balance of energy flow from light absorption into biomass was investigated under simulated natural light conditions in the diatom Phaeodactylum tricornutum and the green alga Chlorella vulgaris. The energy balance was quantified by comparative analysis of carbon accumulation in the new biomass with photosynthetic electron transport rates per absorbed quantum, measured both by fluorescence quenching and oxygen production. The difference between fluorescence- and oxygen-based electron flow is defined as 'alternative electron cycling'. The photosynthetic efficiency of biomass production was found to be identical for both algae under nonfluctuating light conditions. In a fluctuating light regime, a much higher conversion efficiency of photosynthetic energy into biomass was observed in the diatom compared with the green alga. The data clearly show that the diatom utilizes a different strategy in the dissipation of excessively absorbed energy compared with the green alga. Consequently, in a fluctuating light climate, the differences between green algae and diatoms in the efficiency of biomass production per photon absorbed are caused by the different amount of alternative electron cycling.     

Functional integration of the HUP1 hexose symporter gene into the genome of C. reinhardtii: Impacts on biological H(2) production

Phototrophic organisms use photosynthesis to convert solar energy into chemical energy. In nature, the chemical energy is stored in a diverse range of biopolymers. These sunlight-derived, energy-rich biopolymers can be converted into environmentally clean and CO(2) neutral fuels. A select group of photosynthetic microorganisms have developed the ability to extract and divert protons and electrons derived from water to chloroplast hydrogenase(s) to produce molecular H(2) fuel. Here, we describe the development and characterization of C. reinhardtii strains, derived from the high H(2) production mutant Stm6, into which the HUP1 (hexose uptake protein) hexose symporter from Chlorella kessleri was introduced. The isolated cell lines can use externally supplied glucose for heterotrophic growth in the dark. More importantly, external glucose supply (1mM) was shown to increase the H(2) production capacity in strain Stm6Glc4 to approximately 150% of that of the high-H(2) producing strain, Stm6. This establishes the foundations for a new fuel production process in which H(2)O and glucose can simultaneously be used for H(2) production. It also opens new perspectives on future strategies for improving bio-H(2) production efficiency under natural day/night regimes and for using sugar waste material for energy production in green algae as photosynthetic catalysts.     

Regulatory factors for the assembly of thylakoid membrane protein complexes

Major multi-protein photosynthetic complexes, located in thylakoid membranes, are responsible for the capture of light and its conversion into chemical energy in oxygenic photosynthetic organisms. Although the structures and functions of these photosynthetic complexes have been explored, the molecular mechanisms underlying their assembly remain elusive. In this review, we summarize current knowledge of the regulatory components involved in the assembly of thylakoid membrane protein complexes in photosynthetic organisms. Many of the known regulatory factors are conserved between prokaryotes and eukaryotes, whereas others appear to be newly evolved or to have expanded predominantly in eukaryotes. Their specific features and fundamental differences in cyanobacteria, green algae and land plants are discussed.     

Acclimation of green algae to sulfur deficiency: underlying mechanisms and application for hydrogen production

Hydrogen is definitely one of the most acceptable fuels in the future. Some photosynthetic microorganisms, such as green algae and cyanobacteria, can produce hydrogen gas from water by using solar energy. In green algae, hydrogen evolution is coupled to the photosynthetic electron transport in thylakoid membranes via reaction catalyzed by the specific enzyme, (FeFe)-hydrogenase. However, this enzyme is highly sensitive to oxygen and can be quickly inhibited when water splitting is active. A problem of incompatibility between the water splitting and hydrogenase reaction can be overcome by depletion of algal cells of sulfur which is essential element for life. In this review the mechanisms underlying sustained hydrogen photoproduction in sulfur deprived C. reinhardtii and the recent achievements in studying of this process are discussed. The attention is focused on the biophysical and physiological aspects of photosynthetic response to sulfur deficiency in green algae.     

水葫芦对藻类的克制效应

水葫芦对藻类生长有克制作用。其机制主要是由于水葫芦根系向水体分泌的有机物质能伤害和杀死藻类。用水葫芦种植水培养藻类,使藻类的光合作用速率显著降低,叶绿素a破坏,细胞还原TTC的能力下降。在荧光显微镜下可看到藻细胞从鲜红色变为淡蓝绿色。     

微藻光合作用制氢——能源危机的最终出路?

微藻光合作用制氢是解决能源短缺问题的有效途径.本文介绍了微藻光合作用制氢的机理,包括蓝藻固氮酶和可逆氢酶产氢以及绿藻可逆氢酶产氢的机理.在分析光合制氢限制因素的基础上,指出筛选和构建高效放氢藻株是制氢的有效途径.然后介绍了“直接生物光解”、固氮酶放氢和“间接生物光解”等制氢方式.利用绿藻“间接生物光解”水制氢是一种最有发展潜力的制氢方式.本文最后展望了微藻光合制氢的前景.     

The unique features of starch metabolism in red algae.

Red algae (Rhodophyceae) are photosynthetic eukaryotes that accumulate starch granules outside of their plastids. The starch granules from red algae (floridean starch) show structural similarities with higher plant starch granules but lack amylose. Recent studies have indicated that the extra-plastidic starch synthesis in red algae proceeds via a UDP glucose-selective alpha-glucan synthase, in analogy with the cytosolic pathway of glycogen synthesis in other eukaryotes. On the other hand, plastidic starch synthesis in green cells occurs selectively via ADP glucose in analogy with the pathway of glycogen synthesis in prokaryotes from which plastids have evolved. Given the emerging consensus of a monophyletic origin of plastids, it would appear that the capacity for starch synthesis selectively evolved from the alpha-glucan synthesizing machinery of the host ancestor and its endosymbiont in red algae and green algae, respectively. This implies the evolution of fundamentally different functional relationships between the different subcellular compartments with regard to photosynthetic carbon metabolism in these organisms. It is suggested that the biochemical and molecular elucidation of floridean starch synthesis may offer new insights into the metabolic strategies of photosynthetic eukaryotes.     

Tolerance of soil algae and cyanobacteria to drought stress

Tolerance to drought stress in soil crust microorganisms is essential for exploiting suitable organisms for restoring soil. In this study, the responses to drought stress of two drought‐tolerant species, a green alga and a cyanobacterium, were compared with those of two non‐tolerant green algae. In response to drought stress, induced by treatment with polyethylene glycol, the intracellular proline levels increased and were associated with increases in malondialdehye, pigment contents, and enzyme activities such as superoxide dismutase (SOD) and peroxidase (POD). Our results suggest that tolerance to drought stress could be indicated by the intracellular levels of proline, SOD, and carotenoids. This study provides insights into the drought physiology of the photosynthetic microorganisms and suggests that Leptolyngbya boryana and Chlorella vulgaris are suitable pioneer organisms for soil restoration.     

Chlororespiration in Green Algae Isolated From Desert Crusts

Photosynthetic organisms enduring extreme temperatures, low water availability, or high light require photoprotective mechanisms to prevent sustained damage to photosynthetic machinery. Green microalgae living in desert crust communities of the south‐western US experience all these environmental stresses, yet photophysiological studies of green algae in the literature have focused on only a handful of common aquatic and marine species. We are examining the variation in green algal photoprotective mechanisms that is the result of natural selection acting independently in multiple lineages of highly diverse desert green algae (Chlorophyta) within the classes Chlorophyceae and Trebouxiophyceae. We have found that unusually extensive dark reduction of the plastoquinone pool is a prominent photophysiological feature among these desert algae; this reduction may be linked with enhanced chlororespiration. Recently, chlororespiration in higher plants has been linked through mutant analysis to control of the carotenoid synthetic pathway, heat stress, and starch metabolism among other pathways, though the function of chlororespiration remains controversial. Given that green algae and higher plants are monophyletic, analysis of potential chlororespiration in desert green algae may help decipher the evolution of the chlororespiratory process as well as its potential role in photoprotection in desert habitats.     

关于生物制氢

简述了生物制氢发展过程及现今取得的成果。经济发展和人类对能源需求造成了诸如环境污染、常规能源短缺等一系列问题。因此 ,作为一种新型、可再生能源 ,氢能研究已经受到了人们高度重视。与其它制氢方法相比 ,生物制氢有着突出的优点 ,尤其是藻类利用太阳能光解水制氢 ,使人们看到了解决能源问题的希望。     

Origin and evolution of organisms as deduced from 5S ribosomal RNA sequences

A phylogenetic tree of most of the major groups of organisms has been constructed from the 352 5S ribosomal RNA sequences now available. The tree suggests that there are several major groups of eubacteria that diverged during the early stages of their evolution. Metabacteria (= archaebacteria) and eukaryotes separated after the emergence of eubacteria. Among eukaryotes, red algae emerged first; and, later, thraustochytrids (a Proctista group), ascomycetes (yeast), green plants (green algae and land plants), "yellow algae" (brown algae, diatoms, and chrysophyte algae), basidiomycetes (mushrooms and rusts), slime- and water molds, various protozoans, and animals emerged, approximately in that order. Three major types of photosynthetic eukaryotes--i.e., red algae (= Chlorophyll a group), green plants (Chl. a + b group) and yellow algae (Chl. a + c)--are remotely related to one another. Other photosynthetic unicellular protozoans--such as Cyanophora (Chl. a), Euglenophyta (Chl. a + b), Cryptophyta (Chl. a + c), and Dinophyta (Chl. a + c)--seem to have separated shortly after the emergence of the yellow algae.      

Into the deep: new discoveries at the base of the green plant phylogeny

Recent data have provided evidence for an unrecognised ancient lineage of green plants that persists in marine deep-water environments. The green plants are a major group of photosynthetic eukaryotes that have played a prominent role in the global ecosystem for millions of years. A schism early in their evolution gave rise to two major lineages, one of which diversified in the world's oceans and gave rise to a large diversity of marine and freshwater green algae (Chlorophyta) while the other gave rise to a diverse array of freshwater green algae and the land plants (Streptophyta). It is generally believed that the earliest-diverging Chlorophyta were motile planktonic unicellular organisms, but the discovery of an ancient group of deep-water seaweeds has challenged our understanding of the basal branches of the green plant phylogeny. In this review, we discuss current insights into the origin and diversification of the green plant lineage.     

ACCLIMATION OF THE PHOTOSYNTHETIC APPARATUS OF PALMARIA PALMATA (RHODOPHYTA) TO LIGHT QUALITIES THAT PREFERENTIALLY EXCITE PHOTOSYSTEM I OR PHOTOSYSTEM II1

Acclimation of the photosynthetic apparatus to light absorbed primarily by phycobilisomes (which transfer energy predominantly to photosystem II) or absorbed by chlorophyll a (mainly present in the antenna of photosystem I) was studied in the macroalga Palmaria palmata L. In addition, the influence of blue and yellow light, exciting chlorophyll a and phycobilisomes, respectively, ivas investigated. All results were compared to a white light control. Complementary chromatic adaptation in terms of an enhanced ratio of phycoerythrin to phycocyanin under green light conditions was observed. Red light (mainly absorbed by chlorophyll a) and green light (mainly absorbed by phycobilisomes) caused an increase of the antenna system, which was not preferentially excited. Yellow and blue light led to intermediate states comparable to each other and white light. Growth was reduced under all light qualities in comparison to white light, especially under conditions preferably exciting phycobilisomes (green light-adapted algae had a 58% lower growth rate compared to white light-adapted algae). Red and blue light-adapted algae showed maximal photosynthetic capacity with white light excitation and significantly lower values with green light excitation. In contrast, green and yellow light-adapted algae exhibited comparable photosynthetic capacities at all excitation wavelengths. Low-temperature fluorescence emission analysis showed an increase of photosystem II emission in red light-adapted algae and a decrease in green light-adapted algae. A small increase of photosystem I emission teas also found in green light-adapted algae, but this was much less than the photosystem II emission increase observed in red light-adapted algae (both compared to phycobilisome emission). Efficiency of energy transfer from phycobilisomes to photosystem II was higher in red than in green light-adapted algae. The opposite was found for the energy transfer efficiency from phycobilisomes to photosystem I. Zeaxanthin content increased in green and blue light-adapted algae compared to red, white, and yellow light-adapted algae. Results are discussed in comparison to published data on unicellular red algae and cyanobacteria.     

Phylogenetic viewpoints on regulation of light harvesting and electron transport in eukaryotic photosynthetic organisms

The comparative study of photosynthetic regulation in the thylakoid membrane of different phylogenetic groups can yield valuable insights into mechanisms, genetic requirements and redundancy of regulatory processes. This review offers a brief summary on the current understanding of light harvesting and photosynthetic electron transport regulation in different photosynthetic eukaryotes, with a special focus on the comparison between higher plants and unicellular algae of secondary endosymbiotic origin. The foundations of thylakoid structure, light harvesting, reversible protein phosphorylation and PSI-mediated cyclic electron transport are traced not only from green algae to vascular plants but also at the branching point between the “green” and the “red” lineage of photosynthetic organisms. This approach was particularly valuable in revealing processes that (1) are highly conserved between phylogenetic groups, (2) serve a common physiological role but nevertheless originate in divergent genetic backgrounds or (3) are missing in one phylogenetic branch despite their unequivocal importance in another, necessitating a search for alternative regulatory mechanisms and interactions.     

Photosynthetic recovery following desiccation of desert green algae (Chlorophyta) and their aquatic relatives

Recent molecular data suggest that desert green algae have evolved from freshwater ancestors at least 14 times in three major classes (Chlorophyceae, Trebouxiophyceae and Charophyceae), offering a unique opportunity to study the adaptation of photosynthetic organisms to life on land in a comparative phylogenetic framework. We examined the photorecovery of phylogenetically matched desert and aquatic algae after desiccation in darkness and under illumination. Desert algae survived desiccation for at least 4 weeks when dried in darkness, and recovered high levels of photosynthetic quantum yield within 1 h of rehydration in darkness. However, when 4 weeks of desiccation was accompanied by illumination, three of six desert taxa lost their ability to recover quantum yield during rehydration in the dark. Aquatic algae, in contrast, recovered very little during dark rehydration following even just 24 h of desiccation. Re-illuminating rehydrated algae produced a nearly complete recovery of quantum yield in all desert and two of five aquatic taxa. These contrasts provide physiological evidence that desert green algae possess mechanisms for photosynthetic recovery after desiccation distinct from those in aquatic relatives, corroborating molecular evidence that they are not happenstance, short-term visitors from aquatic environments. Photosensitivity during desiccation among desert algae further suggests that they may reside in protected microsites within crusts, and species specificity of photosensitivity suggests that disturbances physically disrupting crusts could lead to shifts or losses of taxonomic diversity within these habitats.     

Characterization of the diversification of phospholipid:diacylglycerol acyltransferases in the green lineage

Triacylglycerols have important physiological roles in photosynthetic organisms, and are widely used as food, feed and industrial materials in our daily life. Phospholipid:diacylglycerol acyltransferase (PDAT) is the pivotal enzyme catalyzing the acyl‐CoA‐independent biosynthesis of triacylglycerols, which is unique in plants, algae and fungi, but not in animals, and has essential functions in plant and algal growth, development and stress responses. Currently, this enzyme has yet to be examined in an evolutionary context at the level of the green lineage. Some fundamental questions remain unanswered, such as how PDATs evolved in photosynthetic organisms and whether the evolution of terrestrial plant PDATs from a lineage of charophyte green algae diverges in enzyme function. As such, we used molecular evolutionary analysis and biochemical assays to address these questions. Our results indicated that PDAT underwent divergent evolution in the green lineage: PDATs exist in a wide range of plants and algae, but not in cyanobacteria. Although PDATs exhibit the conservation of several features, phylogenetic and selection‐pressure analyses revealed that overall they evolved to be highly divergent, driven by different selection constraints. Positive selection, as one major driving force, may have resulted in enzymes with a higher functional importance in land plants than green algae. Further structural and mutagenesis analyses demonstrated that some amino acid sites under positive selection are critically important to PDAT structure and function, and may be central in lecithin:cholesterol acyltransferase family enzymes in general.     

Sensing environmental temperature change through imbalances between energy supply and energy consumption: Redox state of photosystem II

A basic requirement of all photosynthetic organisms is a balance between overall energy supply through temperature-independent photochemical reactions and energy consumption through the temperature-dependent biochemical reactions of photosynthetic electron transport and contiguous metabolic pathways. Since the turnover of photosystem II (PSII) reaction centers is a limiting step in the conversion of light energy into ATP and NADPH, any energy imbalance may be sensed through modulation of the redox state of PSII. This can be estimated in vivo by chlorophyll a fluorescence as changes in the redox state of PSII, or photosystem II excitation pressure, which reflects changes in the redox poise of intersystem electron transport carriers. Through comparisons of photosynthetic adjustment, we show that growth at low temperature mimics growth at high light. We conclude that terrestrial plants, green algae and cyanobacteria do not respond to changes in growth temperature or growth irradiance per se, but rather, respond to changes in the redox state of intersystem electron transport as reflected by changes in PSII excitation pressure, We suggest that this chloroplastic redox sensing mechanism may be an important component for sensing abiotic stresses in general. Thus, in addition to its role in energy transduction, the chloroplast may also be considered a primary sensor of environmental change through a redox sensing/signalling mechanism that acts synergistically with other signal transduction pathways to elicit the appropriate molecular and physiological responses.     

Micro and macroalgal biomass: A renewable source for bioethanol

Population outburst together with increased motorization has led to an overwhelming increase in the demand for fuel. In the milieu of economical and environmental concern, algae capable of accumulating high starch/cellulose can serve as an excellent alternative to food crops for bioethanol production, a green fuel for sustainable future. Certain species of algae can produce ethanol during dark-anaerobic fermentation and thus serve as a direct source for ethanol production. Of late, oleaginous microalgae generate high starch/cellulose biomass waste after oil extraction, which can be hydrolyzed to generate sugary syrup to be used as substrate for ethanol production. Macroalgae are also harnessed as renewable source of biomass intended for ethanol production. Currently there are very few studies on this issue, and intense research is required in future in this area for efficient utilization of algal biomass and their industrial wastes to produce environmentally friendly fuel bioethanol.     

Photobiological hydrogen production: Recent advances and state of the art

Photobiological hydrogen production has advanced significantly in recent years, and on the way to becoming a mature technology. A variety of photosynthetic and non-photosynthetic microorganisms, including unicellular green algae, cyanobacteria, anoxygenic photosynthetic bacteria, obligate anaerobic, and nitrogen-fixing bacteria are endowed with genes and proteins for H₂-production. Enzymes, mechanisms, and the underlying biochemistry may vary among these systems; however, they are all promising catalysts in hydrogen production. Integration of hydrogen production among these organisms and enzymatic systems is a recent concept and a rather interesting development in the field, as it may minimize feedstock utilization and lower the associated costs, while improving yields of hydrogen production. Photobioreactor development and genetic manipulation of the hydrogen-producing microorganisms is also outlined in this review, as these contribute to improvement in the yield of the respective processes.     

The extrinsic proteins of Photosystem II

Years of genetic, biochemical, and structural work have provided a number of insights into the oxygen evolving complex (OEC) of Photosystem II (PSII) for a variety of photosynthetic organisms. However, questions still remain about the functions and interactions among the various subunits that make up the OEC. After a brief introduction to the individual subunits Psb27, PsbP, PsbQ, PsbR, PsbU, and PsbV, a current picture of the OEC as a whole in cyanobacteria, red algae, green algae, and higher plants will be presented. Additionally, the role that these proteins play in the dynamic life cycle of PSII will be discussed.