Academic literature on the topic 'Marine photosynthetic plants'

The low diffusion rates of solutes in water call for a separation of photosynthetic carbon acquirement in aquatic plants into carbon transport and the subsequent photosynthetic reduction of CO2. This paper will focus on the transport of inorganic carbon from the external medium to the site of fixation in marine macrophytes. In accord with the much higher concentration of HCO3− than of CO2 in seawater, most marine macrophytes can utilize the ionic carbon form for their photosynthetic needs. The two known ways of HCO, utilization are (a) via extracellular, carbonic anhydrase catalyzed dehydration of HCO3− to form CO2, which then diffuses into the photosynthesizing cells, and (b) by direct uptake via a transporter. While the first way may be sufficient to support low rates of photosynthesis in temperate regions, it is viewed as futile under situations where high temperatures and irradiances would cause a high pH to form close to the uptake site of carbon and where, consequently, the CO2/HCO3− ratio would be very low. Therefore, it may well be that the direct HCO3− uptake mechanism described for Ulva from more tropical regions confers an adaptational advantage under conditions conducive to higher photosynthetic rates.

Climate change-related events, such as marine heatwaves, are increasing seawater temperatures, thereby putting pressure on marine biota. The cosmopolitan distribution and significant contribution to marine primary production by the genus Ruppia makes them interesting organisms to study thermal tolerance and local adaptation. In this study, we investigated the photosynthetic responses in Ruppia to the predicted future warming in two contrasting bioregions, temperate Sweden and tropical Thailand. Through DNA barcoding, specimens were determined to Ruppia cirrhosa for Sweden and Ruppia maritima for Thailand. Photosynthetic responses were assessed using pulse amplitude-modulated fluorometry, firstly in short time incubations at 18, 23, 28, and 33 °C in the Swedish set-up and 28, 33, 38, and 43 °C in the Thai set-up. Subsequent experiments were conducted to compare the short time effects to longer, five-day incubations in 28 °C for Swedish plants and 40 °C for Thai plants. Swedish R. cirrhosa displayed minor response, while Thai R. maritima was more sensitive to both direct and prolonged temperature stress with a drastic decrease in the photosynthetic parameters leading to mortality. The results indicate that in predicted warming scenarios, Swedish R. cirrhosa may sustain an efficient photosynthesis and potentially outcompete more heat-sensitive species. However, populations of the similar R. maritima in tropical environments may suffer a decline as their productivity will be highly reduced.

Abstract. Some effects in the biosphere from the Total Solar Eclipse of 29 March 2006 were investigated in field crops and marine zooplankton. Taking into account the decisive role of light on plant life and productivity, measurements of photosynthesis and stomatal behaviour were conducted on seven important field-grown cereal and leguminous crops. A drop in photosynthetic rates, by more than a factor of 5 in some cases, was observed, and the minimum values of photosynthetic rates ranged between 3.13 and 10.13 μmol CO2 m−2 s−1. The drop in solar irradiance and the increase in mesophyll CO2-concentration during the eclipse did not induce stomatal closure thus not blocking CO2 uptake by plants. Light effects on the photochemical phase of photosynthesis may be responsible for the observed depression in photosynthetic rates. Field studies addressing the migratory responses of marine zooplankton (micro-zooplankton (ciliates), and meso-zooplankton) due to the rapid changes in underwater light intensity were also performed. The light intensity attenuation was simulated with the use of accurate underwater radiative transfer modeling techniques. Ciliates, responded to the rapid decrease in light intensity during the eclipse adopting night-time behaviour. From the meso-zooplankton assemblage, various vertical migratory behaviours were adopted by different species.

Abstract. The effects in the biosphere from the Total Solar Eclipse of 29 March 2006 were investigated in field crops and marine zooplankton. Taking into account the decisive role of light on the photoenergetic and photoregulatory plant processes, measurements of photosynthesis and stomatal behaviour were conducted on seven important field-grown cereal and leguminous crops. A drop in photosynthetic rates, by more than a factor of 5 in some cases, was observed, and the minimum values of photosynthetic rates ranged between 3.13 and 10.13 μmol CO2 m−2 s−1. However, since solar irradiance attenuation has not at the same time induced stomatal closure thus not blocking CO2 uptake by plants, it is probably other endogenous factors that has been responsible for the observed fall in photosynthetic rates. Field studies addressing the migratory responses of marine zooplankton (micro-zooplankton (ciliates), and meso-zooplankton) due to the rapid changes in underwater light intensity were also performed. The light intensity attenuation was simulated with the use of accurate underwater radiative transfer modeling techniques. Ciliates, responded to the rapid decrease in light intensity during the eclipse adopting night-time behaviour. From the meso-zooplankton assemblage, various vertical migratory behaviours were adopted by different species.

Carotenoids are natural fat-soluble pigments synthesized by plants, algae, fungi and microorganisms. They are responsible for the coloration of different photosynthetic organisms. Although they play a role in photosynthesis, they are also present in non-photosynthetic plant tissues, fungi, and bacteria. These metabolites have mainly been used in food, cosmetics, and the pharmaceutical industry. In addition to their utilization as pigmentation, they have significant therapeutically applications, such as improving immune system and preventing neurodegenerative diseases. Primarily, they have attracted attention due to their antioxidant activity. Several statistical investigations indicated an association between the use of carotenoids in diets and a decreased incidence of cancer types, suggesting the antioxidant properties of these compounds as an important factor in the scope of the studies against oxidative stress. Unusual marine environments are associated with a great chemical diversity, resulting in novel bioactive molecules. Thus, marine organisms may represent an important source of novel biologically active substances for the development of therapeutics. Marine carotenoids (astaxanthin, fucoxanthin, β-carotene, lutein but also the rare siphonaxanthin, sioxanthin, and myxol) have recently shown antioxidant properties in reducing oxidative stress markers. Numerous of bioactive compounds such as marine carotenoids have low stability, are poorly absorbed, and own very limited bioavailability. The new technique is nanoencapsulation, which can be used to preserve marine carotenoids and their original properties during processing, storage, improve their physiochemical properties and increase their health-promoting effects. This review aims to describe the role of marine carotenoids, their potential applications and different types of advanced nanoformulations preventing and treating oxidative stress related disorders.

AbstractGeneration of ion electrochemical potential differences by primary active transport can involve energy inputs from light, from exergonic redox reactions and from exergonic ATP hydrolysis. These electrochemical potential differences are important for homoeostasis, for signalling, and for energizing nutrient influx. The three main ions involved are H+, Na+ (efflux) and Cl− (influx). In prokaryotes, fluxes of all three of these ions are energized by ion-pumping rhodopsins, with one archaeal rhodopsin pumping H+into the cells; among eukaryotes there is also an H+ influx rhodopsin in Acetabularia and (probably) H+ efflux in diatoms. Bacteriochlorophyll-based photoreactions export H+ from the cytosol in some anoxygenic photosynthetic bacteria, but chlorophyll-based photoreactions in marine cyanobacteria do not lead to export of H+. Exergonic redox reactions export H+ and Na+ in photosynthetic bacteria, and possibly H+ in eukaryotic algae. P-type H+- and/or Na+-ATPases occur in almost all of the photosynthetic marine organisms examined. P-type H+-efflux ATPases occur in charophycean marine algae and flowering plants whereas P-type Na+-ATPases predominate in other marine green algae and non-green algae, possibly with H+-ATPases in some cases. An F-type Cl−-ATPase is known to occur in Acetabularia. Some assignments, on the basis of genomic evidence, of P-type ATPases to H+ or Na+ as the pumped ion are inconclusive.

Iron chronically limits aquatic photosynthesis, especially in marine environments, and the correct perception and maintenance of iron homeostasis in photosynthetic bacteria, including cyanobacteria, is therefore of global significance. Multiple adaptive mechanisms, responsive promoters, and posttranscriptional regulators have been identified, which allow cyanobacteria to respond to changing iron concentrations. However, many factors remain unclear, in particular, how iron status is perceived within the cell. Here we describe a cyanobacterial ferredoxin (Fed2), with a unique C-terminal extension, that acts as a player in iron perception. Fed2 homologs are highly conserved in photosynthetic organisms from cyanobacteria to higher plants, and, although they belong to the plant type ferredoxin family of [2Fe-2S] photosynthetic electron carriers, they are not involved in photosynthetic electron transport. As deletion offed2appears lethal, we developed a C-terminal truncation system to attenuate protein function. Disturbed Fed2 function resulted in decreased chlorophyll accumulation, and this was exaggerated in iron-depleted medium, where different truncations led to either exaggerated or weaker responses to low iron. Despite this, iron concentrations remained the same, or were elevated in all truncation mutants. Further analysis established that, when Fed2 function was perturbed, the classical iron limitation marker IsiA failed to accumulate at transcript and protein levels. By contrast, abundance of IsiB, which shares an operon withisiA, was unaffected by loss of Fed2 function, pinpointing the site of Fed2 action in iron perception to the level of posttranscriptional regulation.

Cyanobacteria are important photosynthetic organisms inhabiting a range of dynamic environments. This phylum is distinctive among photosynthetic organisms in containing genes encoding uncharacterized cystathionine β-synthase (CBS)–chloroplast protein (CP12) fusion proteins. These consist of two domains, each recognized as stand-alone photosynthetic regulators with different functions described in cyanobacteria (CP12) and plants (CP12 and CBSX). Here we show that CBS–CP12 fusion proteins are encoded in distinct gene neighborhoods, several unrelated to photosynthesis. Most frequently, CBS–CP12 genes are in a gene cluster with thioredoxin A (TrxA), which is prevalent in bloom-forming, marine symbiotic, and benthic mat cyanobacteria. Focusing on a CBS–CP12 fromMicrocystis aeruginosaPCC 7806 encoded in a gene cluster with TrxA, we reveal that the domain fusion led to the formation of a hexameric protein. We show that the CP12 domain is essential for hexamerization and contains an ordered, previously structurally uncharacterized N-terminal region. We provide evidence that CBS–CP12, while combining properties of both regulatory domains, behaves different from CP12 and plant CBSX. It does not form a ternary complex with phosphoribulokinase (PRK) and glyceraldehyde-3-phosphate dehydrogenase. Instead, CBS–CP12 decreases the activity of PRK in an AMP-dependent manner. We propose that the novel domain architecture and oligomeric state of CBS–CP12 expand its regulatory function beyond those of CP12 in cyanobacteria.

A descendant of the red algal lineage, diatoms are unicellular eukaryotic algae characterized by thylakoid membranes that lack the spatial differentiation of stroma and grana stacks found in green algae and higher plants. While the photophysiology of diatoms has been studied extensively, very little is known about the spatial organization of the multimeric photosynthetic protein complexes within their thylakoid membranes. Here, using cryo-electron tomography, proteomics, and biophysical analyses, we elucidate the macromolecular composition, architecture, and spatial distribution of photosystem II complexes in diatom thylakoid membranes. Structural analyses reveal 2 distinct photosystem II populations: loose clusters of complexes associated with antenna proteins and compact 2D crystalline arrays of dimeric cores. Biophysical measurements reveal only 1 photosystem II functional absorption cross section, suggesting that only the former population is photosynthetically active. The tomographic data indicate that the arrays of photosystem II cores are physically separated from those associated with antenna proteins. We hypothesize that the islands of photosystem cores are repair stations, where photodamaged proteins can be replaced. Our results strongly imply convergent evolution between the red and the green photosynthetic lineages toward spatial segregation of dynamic, functional microdomains of photosystem II supercomplexes.

Aquatic C4 photosynthesis probably arose in response to dissolved CO2 limitations, possibly before its advent in terrestrial plants. Of over 7600 C4 species, only about a dozen aquatic species are identified. Amphibious Eleocharis species (sedges) have C3–C4 photosynthesis and Kranz anatomy in aerial, but not submersed, leaves. Aquatic grasses have aerial and submersed leaves with C4 or C3–C4 photosynthesis and Kranz anatomy, but some lack Kranz anatomy in the submersed leaves. Two freshwater submersed monocots, Hydrilla verticillata and possibly Egeria densa, are C4 NADP-malic enzyme (NADP-ME) species. A marine macroalga, Udotea flabellum (Chlorophyta), and possibly a diatom, are C4, so it is not confined to angiosperms. Submersed C4 species differ from terrestrial in that β-carboxylation is cytosolic with chloroplastic decarboxylation and Rubisco carboxylation, so the C4 and Calvin cycles operate in the same cell without Kranz anatomy. Unlike terrestrial plants, Hydrilla is a facultative C4 that shifts from C3 to C4 in low [CO2]. It is well documented, with C4 gas exchange and pulse-chase characteristics, enzyme kinetics and localization, high internal [CO2], relative growth rate, and quantum yield studies. It has multiple phosphoenolpyruvate carboxylase isoforms with C3-like sequences. Hvpepc4 appears to be the photosynthetic form induced in C4 leaves, but it differs from terrestrial C4 isoforms in lacking a C4 signature Serine. The molecular mass of NADP-ME (72 kDa) also resembles a C3 isoform. Hydrilla belongs to the ancient Hydrocharitaceae family, and gives insight to early C4 development. Hydrilla is an excellent ‘minimalist’ system to study C4 photosynthesis regulation without anatomical complexities.

The cycling of arsenic in the marine photosynthetic plants and algae was examined by analysing total arsenic concentrations and arsenic species in selected marine photosynthetic organisms from the south-east coast, NSW, Australia. A range of elements required for metabolism in photosynthetic organisms were also analysed to determine if any relationship between these elements and arsenic concentrations occurred. Organisms were selected from salt marsh and mangrove ecosystems, marine inter-tidal and estuarine environments, and two species of marine phytoplankton cultured, to represent the different marine environments that primary producers inhabit. Organisms selected were compared to species within their own environment and then a comparison made between the varying ecosystems. In the salt marsh and mangrove ecosystems, the leaves of four species, the mangrove Avicennia marina, the samphire Sarcocornia quinqueflora, the seablight Suaeda australis, and the seagrass Posidonia australis were sampled from three locations from the south-east coast of NSW using nested sampling. Mean total arsenic concentrations (mean � sd) dry mass for all locations were A. marina (0.38 � 0.18 �g g-1 to 1.2 � 0.7 �g g-1), S. quinqueflora (0.13 � 0.06 �g g-1 to 0.46 � 0.22 �g g-1), S. australis (0.03 � 0.06 �g g-1 to 0.05 � 0.03 �g g-1)and P. australis (0.34 � 0.10 �g g-1 to 0.65 � 0.26 �g g-1). Arsenic concentrations were significantly different between species and locations but were consistently low compared to marine macroalgae species. Significant relationships between As and Fe concentrations for A. marina, S. quinqueflora and P. australis and negative relationship between As and Zn concentrations for S. quinqueflora could partially explain arsenic concentrations in these species. No relationship between As and P concentrations were found in this study. All terrestrial species contained predominantly inorganic arsenic in the water extractable and residue fractions with minor concentrations of DMA in the water-soluble fraction. P. australis also contained dimethylated glycerol and phosphate arsenoriboses. The presence of arsenobetaine, arsenocholine and trimethylated glycerol arsonioribose is most likely due to the presence of epiphytes on fronds on P. australis. In contrast, macroalgae contained higher total arsenic concentrations compared to marine terrestrial angiosperms. Total arsenic concentrations also varied between classes of algae: red macroalgae 4.3 �g g-1 to 24.7 �g g-1, green macroalgae 8.0 �g g-1 to 11.0 �g g-1 and blue green algae 10.4 �g g-1 and 18.4 �g g-1. No significant relations were found between As concentrations and concentrations of Fe, Co, Cu, Mn, Mo, Mg, P and Zn concentrations, elements that are required by macroalgae for photosynthesis and growth. Distinct differences between algal classes were found for the proportion of arsenic species present in the lipid and water-soluble fractions, with green algae having a higher proportion of As in lipids than red or estuarine algae. Acid hydrolysis of the lipid extract revealed DMA, glycerol arsenoribose and TMA based arsenolipids. Within water-soluble extracts, red and blue-green algae contained a greater proportion of arsenic as inorganic and simple methylated arsenic species compared to green algae, which contained predominantly glycerol arsenoribose. Arsenobetaine, arsenocholine and tetramethylarsonium was also present in water-soluble extracts but is not normally identified with macroalgae and is again likely due to the presence of attached epiphytes. Residue extracts contained predominantly inorganic arsenic, most likely associated with insoluble constituents of the cell. Mean arsenic concentrations in the green microalgae Dunaliella tertiolecta were 13.3 �g g-1 to 14.5 �g g-1, which is similar to arsenic concentrations found in green macroalgae in this study. Diatom Phaeodactylum tricornutum arsenic concentrations were 1.62 �g g-1 to 2.08 �g g-1. Varying the orthophosphate concentrations had little effect on arsenic uptake of microalgae. D. tertiolecta and P. tricornutum metabolised arsenic, forming simple methylated arsenic species and arsenic riboses. The ratio of phosphate to glycerol arsenoriboses was higher than that normally found in green macroalgae. The hydrolysed lipid fraction contained DMA arsenolipid (16-96%) with minor proportions of phosphate arsenoribose (4-23%). D. tertiolecta at f/10 phosphate concentration, however, contained glycerol arsenoribose and another arsenic lipid with similar retention as TMAO as well as DMA. The similarities between arsenic species in the water-soluble hydrolysed lipids and water-soluble extracts, especially for P. tricornutum, suggests that cells readily bind arsenic within lipids, either for membrane structure or storage, releasing arsenic species into the cytosol as degradation of lipids occurs. Inorganic arsenic was sequestered into insoluble components of the cell. Arsenic species present in D. tertiolecta at lower phosphate concentrations (f/10) were different to other phosphate concentrations (f/2, f/5), and require further investigation to determine whether this is a species-specific response as a result of phosphate deficiency. Although there are similarities in arsenic concentrations and arsenic species in marine photosynthetic organisms, it is evident that response to environmental concentrations of arsenic in uncontaminated environments is dependent on the mode of transfer from the environment, the influence of other elements in arsenic uptake and the ability of the organism to metabolise and sequester inorganic arsenic within the cell. It is not scientifically sound to generalise on arsenic metabolism in �marine plants� when species and the ecosystem in which they exist may influence the transformation of arsenic in higher marine organisms. There is no evidence to suggest that angiosperms produce AB as arsenic is mostly present as inorganic As, with little or no arsenic present in the lipids. However, marine macro- and microalgae both contain lipids with arsenic moieties that may be precursors for AB transformation. Specifically, the presence of TMA and dimethylated arsenoribose based arsenolipids both can transform to AB via intermediates previously identified in marine organisms. Further identification and characterization of As containing lipids is required.

Photosynthesis in the world's oceans by marine algae is responsible for approximately 50% of CO2 fixed into organic carbon. Aquatic primary producers are intricately linked to the climate system due to their reliance on CO2 as a substrate for photosynthesis and role in the removal and export of carbon from the surface ocean to marine sediments. The evolutionary history of marine algae was shaped by changes in the climate system. As a result, fossilized marine algae and modern representatives of ancient groups have the potential to unlock information about the Earth’s climatic past. To use this information and fully understand the role of marine algae in the carbon cycle, however, it is essential to develop an in-depth understanding of CO2 fixation in these organisms. In this thesis I look at carbon fixation in biomineralizing marine algae from a geological and a biological perspective. First I apply a carbon isotope proxy for CO2 to organic material preserved in marine diatom frustules from an extremely transformative period in geological history, the Eocene-Oligocene boundary. Subsequently, this thesis aims to address gaps in our understanding of carbon fixation in eukaryotic marine algae. I present a novel dataset of kinetics of the carbon fixing enzyme, Rubisco, in eukaryotic algae, investigate the role of a pyrenoid-based carbon concentrating mechanism, and identify plastic changes in carbon fixing machinery in response to changing CO2. The findings from this thesis refine our understanding of primary production in the oceans and how we apply algae-based CO2 proxies to understand ancient climates.

The in origin intertidal marine brown alga Fucus vesiculosus L. grow permanently sublittoral in the brackish Bothnian Sea, side by side with the recently discovered F. radicans L. Bergström et L. Kautsky. Environmental conditions like salinity, light and temperature are clearly different between F. vesiculosus growth sites in the Bothnian Sea (4-5 practical salinity units, psu; part of the Baltic Sea) and the tidal Norwegian Sea (34-35 psu; part of the Atlantic Ocean). The general aims of this thesis were to compare physiological aspects between the marine ecotype and the brackish ecotype of F. vesiculosus as well as between the two Bothnian Sea species F. vesiculosus and F. radicans. The result in the study indicates a higher number of water soluble organic compounds in the marine ecotype of F. vesiculosus compared to the brackish ecotype. These compounds are suggested to be compatible solutes and be due to an intertidal and sublittoral adaptation, respectively; where the intertidal ecotype needs the compounds as a protection from oxygen radicals produced during high irradiation at low tide. The sublittoral ecotype might have lost the ability to synthesize these compound/compounds due to its habitat adaptation. The mannitol content is also higher in the marine ecotype compared to the brackish ecotype of F. vesiculosus and this is suggested to be due to both higher level of irradiance and higher salinity at the growth site. 77 K fluorescence emission spectra and immunoblotting of D1 and PsaA proteins indicate that both ecotypes of F. vesiculosus as well as F. radicans have an uneven ratio of photosystem II/photosystem I (PSII/PSI) with an overweight of PSI. The fluorescence emission spectrum of the Bothnian Sea ecotype of F. vesiculosus however, indicates a larger light-harvesting antenna of PSII compared to the marine ecotype of F. vesiculosus and F. radicans. Distinct differences in 77 K fluorescence emission spectra between the Bothnian Sea ecotype of F. vesiculosus and F. radicans confirm that this is a reliable method to use to separate these species. The marine ecotype of F. vesiculosus has a higher photosynthetic maximum (Pmax) compared to the brackish ecotype of F. vesiculosus and F. radicans whereas both the brackish species have similar Pmax. A reason for higher Pmax in the marine ecotype of F. vesiculosus compared to F. radicans is the greater relative amount of ribulose-1.5-bisphosphate carboxylase/oxygenase (Rubisco). The reason for higher Pmax in marine ecotype of F. vesiculosus compare to the brackish ecotype however is not due to the relative amount of Rubisco and further studies of the rate of CO2 fixation by Rubisco is recommended. Treatments of the brackish ecotype of F. vesiculosus in higher salinity than the Bothnian Sea natural water indicate that the most favourable salinity for high Pmax is 10 psu, followed by 20 psu. One part of the explanation to a high Pmax in 10 psu is a greater relative amount of PsaA protein in algae treated in 10 psu. The reason for greater amount of PsaA might be that the algae need to produce more ATP, and are able to have a higher flow of cyclic electron transport around PSI to serve a higher rate of CO2 fixation by Rubisco. However, studies of the rate of CO2 fixation by Rubisco in algae treated in similar salinities as in present study are recommended to confirm this theory.
Fucus vesiculosus L. (Blåstång) är en brunalg som i huvudsak växer i tidvattenzonen i marint vatten men arten klarar också att växa konstant under ytan i det bräckta Bottenhavet. Norska havet och den del av Bottenhavet, där algerna är insamlade i denna studie, har salthalterna 34-35 psu (praktisk salthaltsenhet) respektive 4-5 psu. F. radicans L. Bergström et L. Kautsky (Smaltång) är en nyligen upptäckt art (2005) som har utvecklats i Bottenhavet. F. radicans och Bottenhavets ekotyp av F. vesiculosus växer sida vid sida och har tidigare ansetts vara samma art. Sett till hela Östersjön, så ändras ytans salthalt från 25 till 1-2 psu mellan Östersjöns gräns mot Kattegatt och norra Bottenviken. Den låga salthalten i Östersjön beror på det höga flödet av sötvatten från älvarna och på ett litet inflödet av saltvatten i inloppet vid Kattegatt. Salthaltsgradienten är korrelerad med antalet arter som minskar med minskad salthalt. Östersjön är ett artfattigt hav och de arter som finns är till stor del en blandning av söt- och saltvattenarter. Det finns bara ett fåtal arter som är helt anpassade till bräckt vatten och F. radicans är en av dem. Exempel på miljöskillnader för F. vesiculosus i Norska havet och i Bottenhavet är salthalten, tidvattnet, ljuset och temperaturen. Tidvattnet i Norska havet gör att algerna växlar mellan att vara i vattnet och på land, vilket utsätter algerna för stora ljusskillnader, snabba och stora temperaturväxlingar samt även torka. De alger som växer i Bottenhavet har däremot en jämnare och lägre temperatur, istäcke på vintern och mindre tillgång på ljus eftersom de alltid lever under vattenytan. Skillnaderna i miljön mellan växtplatserna leder till skillnader i fysiologiska anpassningar. Anledningen till att F. vesiculosus och F. radicans valdes som studieobjekt i denna avhandling är att de är viktiga nyckelarter i Bottenhavet. F. vesiculosus och F. radicans är de enda större bältesbildande alger som finns i det artfattiga ekosystemet och de används därför flitigt som mat, gömställe, parningsplats och barnkammare för t.ex. fisk. Att de är nyckelarter gör det angeläget att försöka förstå hur algerna är anpassade och hur de reagerar på miljöförändringar för att få veta hur de kan skyddas och bevaras. F. radicans inkluderades även för att se hur en naturlig art i Bottenhavet är anpassad i jämförelse med den invandrade F. vesiculosus. Marin F. vesiculosus inkluderades för att vara en artreferens från artens naturliga växtplats. Studien visar att det finns fler vattenlösliga organiska substanser (finns vissa organiska substanser som har en proteinskyddande funktion) i den marina ekotypen av of F. vesiculosus än i Bottenhavets ekotyp. Anledningen till detta föreslås vara en anpassning till att växa i tidvattenzonen. Vid lågvatten utsätts F. vesiculosus från Norska havet för starkt ljus, uttorkning, och snabba temperatur- växlingar vilket gör att den kan behöva dessa organiska substanser som skydd mot fria syreradikaler som bildas under lågvattenexponeringarna. F. vesiculosus från Bottenhavet har troligen mist förmågan att syntetisera dessa substanser på grund av anpassning till att hela tiden växa under ytan. Mängden mannitol (socker) är högre i den marina ekotypen av of F. vesiculosus än i Bottenhavets ekotyp. Detta föreslås bero på högre fotosyntetiskt maximum i F. vesiculosus från Norska havet jämfört med ekotypen från Bottenhavet. Skillnaden i fotssyntetiskt maximum är bland annat kopplat till ljus- och salthaltskillnaden på algernas växtplatser. Denna teori styrks av att både fotosyntesen och halten av mannitol ökar i Bottenhavets ekotyp när den behandlas i högre salthalt. Studien visar även att båda ekotyperna av F. vesiculosus samt F. radicans har ett ojämnt förhållande mellan fotosystem II och I (PSII och PSI) med en dominans av PSI. Denna slutsats är baserad på fluorescens emissions mätningar vid 77 K (-196 °C) och mätning av den relativa mängden D1 protein (motsvarar PSII) och PsaA protein (motsvarar PSI). F. vesiculosus från Bottenhavet visar ett emission spektrum som pekar mot en jämnare fördelning av PSII och PSI jämfört med den marina ekotypen och F. radicans. Detta stämmer dock inte med förhållandet mellan D1/PsaA som indikerar att alla tre har mer PSI än PSII. Förklaringen till avvikelsen mellan metoderna antas vara att F. vesiculosus från Bottenhavet har större ljus-infångande antennpigment än marin F. vesiculosus och F. radicans. De tydliga skillnaderna i 77 K fluorescens emission spektra mellan Bottenhavets F. vesiculosus och F. radicans visar att denna metod kan användas som säker artidentifiering. Den marina ekotypen av F. vesiculosus har högre fotosyntetiskt maximum än de båda arterna från Bottenhavet. Mätningar av den relativa mängden av enzymet Rubisco, viktigt för upptaget av koldioxid hos växter och alger, visar att mängden enzym är en sannolik förklaring till skillnaden i fotosyntetiskt maximum mellan den marina ekotypen av F. vesiculosus och F. radicans och detta är troligen en normal artskillnad. Mängden Rubisco kan dock inte förklara skillnaden i fotosyntetiskt maximum mellan de båda ekotyperna av F. vesiculosus. För att undersöka vad skillnaden mellan dessa två beror på så föreslås istället mätningar av Rubisco’s koldioxidfixeringshastighet. Det är en ökning av fotosyntetiskt maximum i Bottenhavets ekotyp av F. vesiculosus när den behandlas i högre salthalt (10, 20 och 35 psu) och det högsta fotosyntetiska maximumet uppmättes i alger som behandlats i 10 psu. Denna ökning beror inte på ökning i den relativa mängden av Rubisco. Ökningen i fotosyntesen speglas dock av en ökning av den relativa mängden PsaA. Detta antas bero på att det behövs mer energi i form av ATP och att en ökning av detta kan ske på grund av att mer PsaA kan driva den cykliska elektrontransporten i fotosyntesreaktionen. Ökat behov av ATP antas bero på en ökning av Rubisco aktiviteten men mätning av aktiviteten krävs för att bekräfta detta.

Now that we have discussed how assemblages of marine soft sediments are structured, we need to consider functional aspects. There are a few main interrelationships that need to be discussed here— inter- and intraspecific competition, feeding and predator–prey interactions, the production of biomass, and the production and delivery of recruiting stages. Other functional aspects, such as the effects of pathogens and parasites and the benefits of association (mutualism, parasitism, symbiosis, etc.) are of less importance in the present discussion. By function we mean the rate processes (i.e. those involving time) that either affect (extrinsic processes) or are inside (intrinsic processes and responses) the organisms that live in sediments. Hence these include primary and secondary production and processes that are mitigated by the organisms that live in sediments, such as nutrient and contaminant fluxes into and out of the sediment. We begin with the historical development of the field since such aspects are often overlooked in these days of electronic searches for references. Functional studies of ecosystems really began with Lindeman´s classic paper (1942) on trophic dynamics. Rather than regarding food merely as particulate matter, Lindeman expressed it in terms of the energy it contained, thereby enabling comparisons to be made between different systems. For example, 1 g of the bivalve Ensis is not equivalent in food value to 1 g of the planktonic copepod Calanus, so the two animals cannot be compared in terms of weight, but they can be compared in terms of the energy units that each gram dry weight contains. The energy unit originally used was the calorie, but this has now been superseded by the joule (J), 1 calorie being equivalent to 4.2 joules. Ensis contains 14 654 J g-1 dry wt and Calanus 30 982 J g-1 dry wt. The basic trophic system is well understood and can be summarized as we showed earlier in Fig. I.8 which gives the links between various trophic levels and the role of competition, organic matter transport, and resource partitioning. In systems fuelled by photosynthesis (so excluding the chemosynthetic deep-sea vent systems), the primary source of energy for any community is sunlight, which is fixed and stored in plant material, which thus constitutes the first trophic level in the ecosystem.

The nature of the relationships between physical and biological processes in the ocean is subtle and complex. Not only do the physical phenomena create a structure, such as a shallow, mixed layer or a front, within which biological processes may proceed, but they also influence the rates of biological processes in many indirect ways. In the ocean, physical phenomena control the distribution of nutrients necessary for phytoplankton photosynthesis. Places with higher kinetic energy have higher concentrations of planktonic organisms, and that makes the whole food web richer (Mann and Lazier 1996). For example, in the midriff region of the Sea of Cortés (Tiburón and Ángel de la Guarda; fig. 1.2), tidal currents are strong, and intense mixing occurs, creating a situation similar to constant upwelling. Thus, primary productivity is high, and this area supports large numbers of sea birds and marine mammals (Maluf 1983). The Gulf of California is a dynamic marginal sea of the Pacific Ocean and has been described as an area of great fertility since the time of early explorers. Gilbert and Allen (1943) described it as fabulously rich in marine life, with waters fairly teeming with multitudes of fish, and to maintain these large numbers, there must be correspondingly huge crops of their ultimate food, the phytoplankton. Topographically the gulf is divided into a series of basins and trenches, deepening to the south and separated from each other by transverse ridges (Shepard 1950; fig. 3.1). Input of nutrients into the gulf from rivers is low and has only local coastal effects. The Baja peninsula has only one, very small river, near 27°N; rivers in mainland Mexico and the Colorado River have dams that divert most of the water for agricultural and urban use. The gulf has three main natural fertilization mechanisms: wind-induced upwelling, tidal mixing, and thermohaline circulation. Upwelling occurs off the eastern coast with northwesterly winds (winter conditions from December through May) and off the Baja California coast with southeasterly winds (summer conditions from July through October), with June and November as transition periods (Álvarez-Borrego and Lara- Lara 1991).