The marine environment is an important habitat where many processes occur that affect life conditions on earth. Macrophytes and planktonic oxygen evolvers are an essential component for almost all marine life forms and have developed in an environment that differs largely from the terrestrial habitats. For instance in regards to available ionic forms of inorganic carbon and moving water masses which affects incoming light. It is therefore relevant to examine the physiology of algae and marine plants to identify their unique features and differences to terrestrial plants that once orginated from algae. By using chlorophyll fluorescence measurements alone or combined with measurements of oxygen evolution and protein analysis photosynthetic strategies to withstand excess energy have been evaluated under a variety of experimental conditions. Furthermore metabolic pathways involved in energy transfer from photosynthesis to the site of active carbon uptake have been examined. The following was found: * The ratio between photosynthetic gross oxygen evolution and estimated electron transport rate varies in Ulva spp depending on previous history of light and dark exposures. To obtain P/I curves with ratios close to the theoretical 1:4 value, measurements should be performed on separate pieces of tissue at each irradiance level. * Under carbon deficient conditions, the estimated ETR is larger than the gross oxygen evolution, which may be due to the so called “water-water” cycle and absorption changes in PSII which are not corrected for in the calculation of ETR. * Upon exposure to high irradiances (1500 µmol photons m-2s-1) the PSII core protein D1 is broken down with a concomittant reduction in ETR in Ulva spp. With the decrease in electron transport between PSII and PSI the acidification of the lumen decreases and the ability to dissipate excess energy as heat. At prolonged irradiance, an acclimation occurs with a lesser or no breakdown of D1 indicating an additional photo-protective strategy other than heat dissipation. * Laminaria saccharina is dependent on mitochondrial respiration for active utilization of bicarbonate. By extruding protons outside the plasmalemma an acidification takes place that favors the conversion of bicarbonate into carbon dioxide that then can diffuse in to the cell. These proton pumps are driven by ATP supplied to a large degree from mitochondria, likely through the reductant NADPH produced photochemically. * The marine angiosperm Zostera marina is dependent on mitochondrial respiration for utilization of bicarbonate in a manner similar to that in Laminaria saccharina . However, the water-water cycle may supply additional ATP to the proton pumps in Zostera marina. Both species exhibit a lag-phase at the onset of illumination after a dark incubation period and at least part of this lag-phase is due to a lag in an activation of mitochondrial supported bicarbonate utilization. It is clear that the marine environment holds complex plant and algae species and much is still to discover about the oxygen evolvers that grow beneath the water surface.

The green algae of the genus Halimeda Lamouroux (Chlorophyta, Bryopsidales) and the encrusting loose-lying red coralline algae (Rhodophyta, Corallinales) known as rhodoliths are abundant and widespread in all oceans. They significantly contribute to primary productivity while alive and production of CaCO3 rich sediment materials on death and decay. Carbonate rich sediments are important components in the formation of Coral Reefs and as sources of inorganic carbon (influx) in tropical and subtropical marine environments. This study was initiated to attempt to assess their ecological significance with regard to the above mentioned roles in a tropical lagoon system, Chwaka bay (Indian Ocean), and to address some specific objectives on the genus Halimeda (Chlorophyta, Bryopsidales) and the loose-lying coralline algae (rhodoliths). Four Halimeda species were taxonomically identified in the area. The species identified are the most common inhabitants of the world’s tropical and subtropical marine environments, and no new species were encountered. Using Satellite remote sensing technique in combination with the percentage cover data obtained from ground-truthing field work conducted in the area using quadrants, the spatial and seasonal changes of Submerged Aquatic Macrophytes (SAV) were evaluated. SAV percentage cover through ground-truthing was; 24.4% seagrass, 16% mixed Halimeda spp., 5.3% other macroalgae species while 54.3% remained unvegetated. No significant changes in SAV cover was observed for the period investigated, except in some smaller regions where both loss and gains occurred. The structural complexity of SAV (shoot density, above-ground biomass and canopy height) for most common seagrass communities from six meadows, dominated by Thalassia hemprichii, Enhalus acoroides and Thalassodendron ciliatum, as well as mixed meadows, were estimated and evaluated. Relative growth of Halimeda species was up to 1 segment tip-1 day-1. The number of segments produced was highest in hot season. Differences between the numbers of segments produced were insignificant between the two sites investigated. The C/N ratios obtained probably shows that Halimeda species experience nitrogen limitation in the area and may be a factor among others responsible for the varying growth of species obtained. However, this can be a normal ratio for calcified algae due to high CaCO3 content in their tissues. Standing biomass of mixed Halimeda species averaged between 500-600 g dw m-2 over the bay, while the mean cover in Halimeda meadows was about 1560 g dw m-2. Carbonate production in Halimeda beds varied between 17-57 g CaCO3 m-2 day-1 and for H. macroloba between 12-91 g CaCO3 m-2 day-1. This indicates a high annual input of carbonate in the area. Decomposition of Halimeda using litter bag experiments at site I and II gave a decomposition rate (k) of 0.0064 and k = 0.0091 day-1 ash-free dry weight (AFDW) respectively. Hence it would take 76-103 days for 50% of the materials to decompose. Adding inhibitors or varying the pH significantly reduced inorganic carbon uptake, and demonstrated that the two photosynthesis and calcification were linked. Addition of TRIS strongly inhibited photosynthesis but not calcification, suggesting the involvement of proton pumps in the localized low pH acid zones and high pH basic zones. The high pH zones were maintained by the proton pumps maintaining high calcification, while TRIS was competing for proton uptake from acid zones causing photosynthesis to drop. Rhodoliths were found to maintain high productivity at a temperature of 34oC, and even at 37oC. It is therefore concluded that, rhodoliths are well adapted to high temperatures and excess light, a behaviour which enables them to thrive even in intertidal areas.

Cyanobacteria are an ancient and globally distributed group of photosynthetic prokaryotes including species capable of fixing atmospheric dinitrogen (N2) into biologically available ammonia via the enzyme complex nitrogenase. The ability to form symbiotic interactions with eukaryotic hosts is a notable feature of cyanobacteria and one which, via an ancient endosymbiotic event, led to the evolution of chloroplasts and eventually to the plant dominated biosphere of the globe. Some cyanobacteria are still symbiotically competent and form symbiotic associations with eukaryotes ranging from unicellular organisms to complex plants. Among contemporary plant-cyanobacteria associations, the symbiosis formed between the small fast-growing aquatic fern Azolla and its cyanobacterial symbiont (cyanobiont), harboured in specialized cavities in each Azolla leaf, is the only one which is perpetual and in which the cyanobiont has lost its free-living capacity, suggesting a long-lasting co-evolution between the two partners. In this study, the genome of the cyanobiont in Azolla filiculoides was sequenced to completion and analysed. The results revealed that the genome is in an eroding state, evidenced by a high proportion of pseudogenes and transposable elements. Loss of function was most predominant in genetic categories related to uptake and metabolism of nutrients, response to environmental stimuli and in the DNA maintenance machinery. Conversely, function was retained in key symbiotic processes such as nitrogen-fixation and cell differentiation. A comparative analysis shows that the size of the cyanobiont genome has remained relatively stable, and that few genes have been completely eliminated, since the symbiotic establishment. Indications of genes acquired via horizontal gene transfer were discovered in thec yanobiont genome, some of which may have originated from the bacterial community in the Azolla leaf-cavities. It is concluded that the perpetual nature of the Azolla symbiosis has resulted in pronounced ongoing streamlining of the cyanobiont genome around core symbiotic functions, a process not described previously for complex cyanobacteria or for any bacterial plant symbiont. Further, the status of the genome indicates that the cyanobiont is at an early stage of adapting to its host-restricted environment and continued co-evolution with the plant may result in additional genome reductions. However, although a vertical transmission process is already established, the unusual extracellular location of the cyanobiont and the intricate nature of the symbiosis, may still impose restrictions on such a reductive process. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Submitted. Paper 3: Manuscript.</p>

In the present work the regulation of environmentally induced cell death and signaling of systemic acquired acclimation (SAA) in Arabidopsis thaliana is characterized. We used the lesion simulating disease1 (lsd1) mutant as a model system that is deregulated in light acclimation and programmed cell death (PCD). In this system we identify that redox status controlling SAA and cell death is controlled by the genes LSD1, EDS1, EIN2 and PAD4 which regulate cellular homeostasis of salicylic acid (SA), ethylene (ET), auxin (IAA) and reactive oxygen species (ROS). Furthermore we propose that the roles of LSD1 in light acclimation and in biotic stress are functionally linked. The influence of SA on plant growth, short-term acclimation to high light (HL), and on the redox homeostasis of Arabidopsis leaves was also assessed. SA impaired acclimation of wild-type plants to prolonged conditions of excess excitation energy (EEE). This indicates an essential role of SA in acclimation and regulation of cellular redox homeostasis. We also show that cell death in response to EEE is controlled by specific redox changes of photosynthetic electron transport carriers that normally regulate EEE acclimation. These redox changes cause production of ET that signals through the EIN2 gene and regulon. In the lsd1 mutant, we found that propagation of cell death depends on the plant defence regulators EDS1 and PAD4 operating upstream of ET production. We conclude that the balanced activities of LSD1, EDS1, PAD4 and EIN2 regulate chloroplast dependent acclimatory and defence responses. Furthermore, we show that Arabidopsis hypocotyls form lysigenous aerenchyma in response to hypoxia and that this process involves H2O2 and ET signalling. We found that formation of lysigenous aerenchyma depends on LSD1, EDS1 and PAD4. Conclusively we show that LSD1, EDS1 and PAD4, in their functions as major plant redox and hormone regulators provide a basis for fundamental plant survival in natural contitions.

<p>During photosynthesis, two multi-protein complexes, photosystems (PS) I and II work in tandem to convert the light-energy absorbed by the light-harvesting antennae into chemical energy, which is subsequently used to assimilate atmospheric carbon dioxide into organic carbon compounds. This is the main nutritional basis for life on Earth.</p><p>The photosynthetic antenna of higher plants comprises at least ten different pigment-binding proteins (LHC), which play important roles in photosynthesis. Chlorophyll and carotenoid molecules associated with the LHC proteins are organised into an array, which can be modulated, thereby optimising light-harvesting processes and protection against oxidative damage under conditions of excessive light absorption. All ten LHC proteins have been conserved through eons of evolution, suggesting that there are strong evolutionary pressures to retain all ten proteins, and hence that each protein has a unique function.</p><p>The light-harvesting antenna of higher plant PSI consists of at least four proteins, Lhca1-4, collectively called LHCI. By constructing transgenic Arabidopsis thaliana plants in which each Lhca gene has been individually repressed or knocked-out, a collection of plants with different Lhca protein contents was obtained. The objective was to use these plants to study the structure, function and regulation of the Lhca proteins in vivo. The major findings of this work are as follows.</p><p>Removing single Lhca proteins influenced the stability of the other Lhca proteins, showing that there is a high degree of inter-dependency between the polypeptides in LHCI, and hence that a full set of Lhca proteins is important for maintaining the structural integrity of LHCI. This has provided insight into the organisation of LHCI by revealing clues about the relative positions of each Lhca protein in the antenna complex. The physiological consequences of removing individual Lhca proteins were dependent on the degree of antenna depletion. Plants with relatively small antenna changes could compensate, to some extent, for the loss of LHCI, while larger depletions had profound effects on whole plant resulting in growth reductions.</p><p>The fitness of each Lhca plant was assessed by measuring their seed production in the harsh conditions in the field. We found that all Lhca-deficient plants produced fewer seeds under some conditions, with seed-production compared to wild type varying between 10-80% depending on the extent of LHCI reduction. Therefore, we conclude that each Lhca protein is important for plant fitness, and hence for the survival of the species.</p><p>PSI is characterised by a pool of pigments absorbing light in the red end of the solar visible spectrum, thought to be especially important for plants in dense vegetation systems where the incident light is enriched in wavelengths higher than 690 nm. A majority of these pigments are situated on LHCI and, based on in-vitro studies, were thought to be mainly associated with Lhca4. Using our plants, we have established that red pigments are indeed present on all Lhca proteins and that these pigments become even more red upon association with PSI.</p>

During photosynthesis, two multi-protein complexes, photosystems (PS) I and II work in tandem to convert the light-energy absorbed by the light-harvesting antennae into chemical energy, which is subsequently used to assimilate atmospheric carbon dioxide into organic carbon compounds. This is the main nutritional basis for life on Earth. The photosynthetic antenna of higher plants comprises at least ten different pigment-binding proteins (LHC), which play important roles in photosynthesis. Chlorophyll and carotenoid molecules associated with the LHC proteins are organised into an array, which can be modulated, thereby optimising light-harvesting processes and protection against oxidative damage under conditions of excessive light absorption. All ten LHC proteins have been conserved through eons of evolution, suggesting that there are strong evolutionary pressures to retain all ten proteins, and hence that each protein has a unique function. The light-harvesting antenna of higher plant PSI consists of at least four proteins, Lhca1-4, collectively called LHCI. By constructing transgenic Arabidopsis thaliana plants in which each Lhca gene has been individually repressed or knocked-out, a collection of plants with different Lhca protein contents was obtained. The objective was to use these plants to study the structure, function and regulation of the Lhca proteins in vivo. The major findings of this work are as follows. Removing single Lhca proteins influenced the stability of the other Lhca proteins, showing that there is a high degree of inter-dependency between the polypeptides in LHCI, and hence that a full set of Lhca proteins is important for maintaining the structural integrity of LHCI. This has provided insight into the organisation of LHCI by revealing clues about the relative positions of each Lhca protein in the antenna complex. The physiological consequences of removing individual Lhca proteins were dependent on the degree of antenna depletion. Plants with relatively small antenna changes could compensate, to some extent, for the loss of LHCI, while larger depletions had profound effects on whole plant resulting in growth reductions. The fitness of each Lhca plant was assessed by measuring their seed production in the harsh conditions in the field. We found that all Lhca-deficient plants produced fewer seeds under some conditions, with seed-production compared to wild type varying between 10-80% depending on the extent of LHCI reduction. Therefore, we conclude that each Lhca protein is important for plant fitness, and hence for the survival of the species. PSI is characterised by a pool of pigments absorbing light in the red end of the solar visible spectrum, thought to be especially important for plants in dense vegetation systems where the incident light is enriched in wavelengths higher than 690 nm. A majority of these pigments are situated on LHCI and, based on in-vitro studies, were thought to be mainly associated with Lhca4. Using our plants, we have established that red pigments are indeed present on all Lhca proteins and that these pigments become even more red upon association with PSI.

<p>Mine tailings can be rich in sulphide minerals and may form acid mine drainage (AMD) through reaction with atmospheric oxygen and water. AMD contains elevated levels of metals and arsenic (As) that could be harmful to animals and plants. An oxygen-consuming layer of organic material and plants on top of water-covered tailings would probably reduce oxygen penetration into the tailings and thus reduce the formation of AMD. However, wetland plants have the ability to release oxygen through the roots and could thereby increase the solubility of metals and As. These elements are released into the drainage water, taken up and accumulated in the plant roots, or translocated to the shoots. </p><p>The aim was to examine the effects of plant establishment on water-covered mine tailings by answering following questions: A) Is plant establishment on water-covered mine tailings possible? B) What are the metal and As uptake and translocation properties of these plants? C) How do plants affect metal and As release from mine tailings, and which are the mechanisms involved?</p><p><i>Carex rostrata Stokes, Eriophorum angustifolium</i> Honck., <i>E. scheuchzeri</i> Hoppe, <i>Phragmites australis</i> (Cav.) Steud., <i>Salix phylicifolia</i> L. and <i>S. borealis</i> Fr. were used as test plants. Influences of plants on the release of As, Cd, Cu, Pb, Zn and in some cases Fe in the drainage water, and plant element uptake were studied in greenhouse experiments and in the field. </p><p>The results obtained demonstrate that plant establishment are possible on water-covered unweathered mine tailings, and a suitable amendment was found to be sewage sludge. On acidic, weathered tailings, a pH increasing substance such as ashes should be added to improve plant establishment. The metal and As concentrations of the plant tissue were found to be generally higher in roots than in shoots. The uptake was dependent on the metal and As concentrations of the tailings and the release of organic acids from plant roots may have influenced the uptake. The metal release from tailings into the drainage water caused by<i> E. angustifolium </i>was found to depend greatly on the age and chemical properties of the tailings. However, no effects of <i>E. angustifolium </i>on As release was found. Water from old sulphide-, metal- and As-rich tailings with low buffering capacity were positively affected by <i>E. angustifolium </i>by causing higher pH and lower metal concentrations. In tailings with relatively low sulphide, metal and As contents combined with a low buffering capacity, plants had the opposite impact, i.e. a reduction in pH and elevated metal levels of the drainage water. The total release of metal and As from the tailings, i.e. drainage water together with the contents in shoots and roots, was found to be similar for <i>C. rostrata</i>, <i>E. angustifolium </i>and <i>P. australis</i>, except for Fe and As, where the release was highest for <i>P. australis</i>. The differences in metal and As release from mine tailings were mainly found to be due to the release of O₂from the roots, which changes the redox potential. Release of organic acids from the roots slightly decreased the pH, although did not have any particular influence on the release of metal and As. </p><p>In conclusion, as shown here, phytostabilisation may be a successful technique for remediation of mine tailings with high element and sulphide levels, and low buffering capacity.</p>

<p>Mercury (Hg) pollution is a global environmental problem. Numerous Hg-contaminated sites exist in the world and new techniques for remediation are urgently needed. Phytoremediation, use of plants to remove pollutants from the environment or to render them harmless, is considered as an environment-friendly method to remediate contaminated soil <i>in-situ</i> and has been applied for some other heavy metals. Whether this approach is suitable for remediation of Hg-contaminated soil is, however, an open question. The aim of this thesis was to study the fate of Hg in terrestrial plants (particularly the high biomass producing willow, <i>Salix spp</i>.) and thus to clarify the potential use of plants to remediate Hg-contaminated soils.</p><p>Plants used for phytoremediation of Hg must tolerate Hg. A large variation (up to 30-fold difference) was detected among the six investigated clones of willow in their sensitivity to Hg as reflected in their empirical toxicity threshold (TT_95b), the maximum unit toxicity (UTmax) and EC50 levels. This gives us a possibility to select Hg-tolerant willow clones to successfully grow in Hgcontaminated soils for phytoremediation.</p><p>Release of Hg into air by plants is a concern when using phytoremediation in practice. No evidence was found in this study that Hg was released to the air via shoots of willow, garden pea (<i>Pisum sativum</i> L. cv Faenomen), spring wheat (<i>Triticum aestivum</i> L. cv Dragon), sugar beet (<i>Beta vulgaris </i>L. cv Monohill), oil-seed rape (<i>Brassica napus</i> L. cv Paroll) and white clover (<i>Trifolium repens</i> L.). Thus, we conclude that the Hg burden to the atmosphere via phytoremediation is not increased.</p><p>Phytoremediation processes are based on the ability of plant roots to accumulate Hg and to translocate it to the shoots. Willow roots were shown to be able to efficiently accumulate Hg in hydroponics, however, no variation in the ability to accumulate was found among the eight willow clones using CVAAS to analyze Hg content in plants. The majority of the Hg accumulated remained in the roots and only 0.5-0.6% of the Hg accumulation was translocated to the shoots. Similar results were found for the five common cultivated plant species mentioned above. Moreover, the accumulation of Hg in willow was higher when being cultivated in methyl-Hg solution than in inorganic Hg solution, whereas the translocation of Hg to the shoots did not differ.</p><p>The low bioavailability of Hg in contaminated soil is a restricting factor for the phytoextraction of Hg. A selected tolerant willow clone was used to study whether iodide addition could increase the plant-accumulation of Hg from contaminated soil. Both pot tests and field trials were carried out. Potassium iodide (KI) addition was found to mobilize Hg in contaminated soil and thus increase the bioavailability of Hg in soils. Addition of KI (0.2–1 mM) increased the Hg concentrations up to about 5, 3 and 8 times in the leaves, branches and roots, respectively. However, too high concentrations of KI were toxic to plants. As the majority of the Hg accumulated in the roots, it might be unrealistic to use willow for phytoextraction of Hg in practice, even though iodide could enhance the phytoextraction efficiency.</p><p>In order to study the effect of willow on various soil fractions of Hg-contaminated soil, a 5-step sequential soil extraction method was used. Both the largest Hg-contaminated fractions, i.e. the Hg bound to residual organic matter (53%) and sulphides (43%), and the residual fraction (2.5%), were found to remain stable during cultivations of willow. The exchangeable Hg (0.1%) and the Hg bound to humic and fulvic acids (1.1%) decreased in the rhizospheric soil, whereas the plant accumulation of Hg increased with the cultivation time. The sum of the decrease of the two Hg fractions in soils was approximately equal to the amount of the Hg accumulated in plants. Consequently, plants may be suitable for phytostabilization of aged Hg-contaminated soil, in which root systems trap the bioavailable Hg and reduce the leakage of Hg from contaminated soils.</p>

<p>Although low temperature plays an important role in determining agricultural yield, little is known about the effect on the underlying biochemical and physiological processes that influence plant growth. Photosynthesis and respiration are central to plant growth and both processes are heavily affected by temperature. However, many plants have the ability to cope with low temperature and resume growth by cold acclimating.</p><p>We have shown that enhancement of carbon fixation, an increased flux of carbon into sucrose and the recovery of diurnal export is crucial for the recovery of functional carbon metabolism at low temperature in Arabidopsis thaliana. The recovery of efflux is governed by increased expression of sucrose transporters along with changes in vascularisation. We also demonstrate the importance of controlling the flux of metabolites between the chloroplast and the cytosol by regulating the expression of AtTPT.</p><p>We further investigated the difference in response between leaves developed at low temperature but originating from warm grown Arabidopsis and leaves from plants grown from seed at low temperature. We were able to distinguish factors that respond specifically to low temperature from those that are connected to the actual stress. Substantial difference could be seen in the different metabolomes. One conclusion drawn is that the increase in sucrose reported at low temperature is an essential feature for life in the cold. </p><p>In an extended study we were able to transfer some of the key factor of cold acclimation in Arabidopsis to other species. The study included forbs, grasses and evergreen trees/shrubs showed that there are striking similarities in the extent and biochemical changes that underpin acclimation among the different functional groups.</p><p>Low temperature does not only influence growth of the leaves, perennial organs such as the corm of the ornamental plant Crocus vernus is also affected. However in these plants low temperature has a positive effect on the final size of the corm. We were able to show that this enhanced growth was an affect of increased cell size and thus increased sink capacity, which ultimately delays leaf senescence</p>

<p>The solar energy accumulated by photosynthesis over billions of years is the sole source of energy available on Earth. Photosystem II (PSII) uses the sunlight to split water, an energetically unfavorable reaction where electrons and protons are extracted from water and oxygen is released as a by-product. Understanding this process is crucial for the future development of clean, renewable and unlimited energy sources, which can use sunlight to split water and produce hydrogen and electricity. In order to do so we need to understand how this is solved in plants.</p><p>I have been focusing on the role of two lumenal proteins associated with the thylakoid membrane PsbO and Cah3, in the water oxidation process. Convincing evidences have been presented supporting the hypothesis that bicarbonate acts as a proton acceptor in the water splitting process in PSII and the lumenal carbonic anhydrase, Cah3, supplies bicarbonate required for this function. The PsbO protein, an important constituent of the water-oxidizing complex, however, its function is still unknown. The PsbO protein undergoes a pH dependent conformational change that in turn influences its capacity to bind calcium and manganese, forming a catalytic Mn4Ca cluster in PSII. We propose that light-induced structural dynamics of the PsbO is of functional relevance for the regulation of proton release and for forming a proton sensing - proton transporting pathway. The cluster of conserved glutamic and aspartic acid residues in the PsbO protein acts as buffering antennae providing efficient acceptors of protons derived from substrate water molecules. Both proteins, Cah3 and PsbO have a conserved S-S bridge, required for proper folding and activity; therefore they are potential targets for red-ox regulation in lumen.</p> / <p>Solenergi som omvandlats av fotosyntesen under miljarder av år är basen för nästan all energi på jorden. Fotosystem 2 använder solljuset till att oxidera vatten, ur energisynpunkt en ofördelaktig process, där elektroner och protoner extraheras från vattenmolekyler vilket ger upphov till syrgas som biprodukt. Förståelsen av denna process är viktig för att vi i framtiden skall kunna utveckla rena och förnyelsebara energislag i obegrensad mängd. Genom att efterlikna fotosyntesprocessen skulle vi i framtiden kunna utvecka artificiella system som använder solljuset till att sönderdela vatten för att producera vätgas eller elektrisitet. För att kunna göra det så måste vi kunna förstå hur dessa processer fungerar i växterna.</p><p>Min forskning har fokuserat på att förstå funktionen hos två av de proteiner, PsbO och Cah3, som deltar i sönderdelningen av vatten. Jag har visat, för första gången, att ett lumen karboanhydras, Cah3, deltar i regleringen av den process där vatten spjälkas. Jag postulerar att Cah3 underlättar bort transporten av protoner från det vattenoxiderande komplexet genom att generera bikarbonat lokalt, som kan fungera som proton transportör. PsbO proteinet genomgår en pH beroende konformationsförändring vilket påverkar dess kapacitet and binda calcium och mangan som i sin tur formar ett katalytiskt Mn4Ca center i fotosystem 2. Jag föreslår att en ljusberoende strukturförändring av Psbo är av funktionell betydelse för regleringen av protonfrigörandet och formar ett proton-avkännande och proton-transporterande system. Ett kluster av konserverande glutamat- och aspartat-aminosyror i PsbO proteinet fungerar som ett buffrande nätverk för protoner som frigörs vid oxidering av vatten. Båda dessa proteiner innerhåller S-S bryggor ock kan därför vara red-ox reglerade i lumen.</p>