Oxygen and the light–dark cycle of nitrogenase activity in two unicellular cyanobacteria

Cyanobacteria capable of fixing dinitrogen exhibit various strategies to protect nitrogenase from inactivation by oxygen. The marine Crocosphaera watsonii WH8501 and the terrestrial Gloeothece sp. PCC6909 are unicellular diazotrophic cyanobacteria that are capable of aerobic nitrogen fixation. These cyanobacteria separate the incompatible processes of oxygenic photosynthesis and nitrogen fixation temporally, confining the latter to the dark. Although these cyanobacteria thrive in fully aerobic environments and can be cultivated diazotrophically under aerobic conditions, the effect of oxygen is not precisely known due to methodological limitations. Here we report the characteristics of nitrogenase activity with respect to well‐defined levels of oxygen to which the organisms are exposed, using an online and near real‐time acetylene reduction assay combined with sensitive laser‐based photoacoustic ethylene detection. The cultures were grown under an alternating 12–12 h light–dark cycle and acetylene reduction was recorded continuously. Acetylene reduction was assayed at 20%, 15%, 10%, 7.5%, 5% and 0% oxygen and at photon flux densities of 30 and 76 μmol m^?2 s^?1 provided at the same light–dark cycle as during cultivation. Nitrogenase activity was predominantly but not exclusively confined to the dark. At 0% oxygen nitrogenase activity in Gloeothece sp. was not detected during the dark and was shifted completely to the light period, while C. watsonii did not exhibit nitrogenase activity at all. Oxygen concentrations of 15% and higher did not support nitrogenase activity in either of the two cyanobacteria. The highest nitrogenase activities were at 5–7.5% oxygen. The highest nitrogenase activities in C. watsonii and Gloeothece sp. were observed at 29°C. At 31°C and above, nitrogenase activity was not detected in C. watsonii while the same was the case at 41°C and above in Gloeothece sp. The differences in the behaviour of nitrogenase activity in these cyanobacteria are discussed with respect to their presumed physiological strategies to protect nitrogenase from oxygen inactivation and to the environment in which they thrive.     

Thermal acclimation and photoacclimation of photosynthesis in the brown alga Laminaria saccharina

Thermal acclimation and photoacclimation of photosynthesis were compared in Laminaria saccharina sporophytes grown at temperatures of 5 and 17 °C and irradiances of 15 and 150μmol photons m^?2 s^?1. When measured at a standard temperature (17°C), rates of light-saturated photosynthesis (P_max) were higher in 5 °C-grown algae (c. 3.0 μmol O₂ m^?2 s^?1) than in 17 °C-grown algae (c. 0.9 μmol O₂ m_-2 s_-1). Concentrations of Rubisco were also 3-fold higher (per unit protein) in 5 °C-grown algae than in algae grown at 17 °C. Light-limited photosynthesis responded similarly to high temperature and low light Photon yields (α) were higher in algae grown at high temperature (regardless of light), and at 5 °C in low light, than in algae grown at 5 °C in high light Differences in a were correlated with light absorption; both groups of 17 °C algae and 5 °C low-light algae absorbed c. 75% of incident light, whereas 5 °C high-light algae absorbed c. 55%. Increased absorption was correlated with increases in pigment content PSII reaction centre densities and the fucoxanthin-Chl ale protein complex (FCP). Changes in a were also attributed, in part, to changes in the maximum photon yield of photosynthesis (0_max). PSI reaction centre densities were unaffected by growth temperature, but the areal concentration of PSI in low-light-grown algae was twice that of high-light-grown algae (c. 160.0 versus 80.0 nmol m^?2). We suggest that complex metabolic regulation allows L, saccharina to optimize photosynthesis over the wide range of temperatures and light levels encountered in nature.     

Nitrogenase Activity in the Lichen Stereocaulon paschale: Recovery after Dry Storage

The effect of prolonged storage on nitrogenase activity in Stereocaulon paschale (L.) Fr. was studied. The thalli had a very low water content during the storage over silica gel at 4°C in the dark. For C₂H₂-reduction measurements the lichen samples were remoistened and held at 15°C in the light for 1–36 h before the addition of C₂H₂. With the longest pre-incubation time preceding the nitrogenase activity measurements there was no significant decrease in nitrogenase activity during 75 weeks of storage. The method of storage was simple and inexpensive and the nitrogenase activity characteristic of each lichen batch, due to seasonal variation, was well preserved during the storage.     

The Adaptation of Plankton Algae

The various aspects of the adaptation of plankton algae lo light and temperature are discussed. The shape of a light intensity-photosynthesis curve is shown to be an important means of describing the physiological adjustment of an algal population. If the algae are not exposed to adverse influences such as poisons, pronounced nutrient deficiency or light shocks, the rate of real photosynthesis per mg chlorophyll a at 1 Klux (incandescent light) should be about 0.4–0.6 mg C/hour. Hence this rate presents an excellent means of judging the quality of experiments. Experiments are presented where Chlorella pyrenoidosa was adapted to light intensities between 0.32 klux and 21 Klux. This alga adapts to different light intensities by varying the amount of pigments per cell. Algae grown at 1 Klux have about 10 times more chlorophyll per cell than those grown at 21 klux. Other species of algae—but by no means all—are shown to behave in the same way. The problem of algal resistance to photo-oxidation at high light intensities is discussed. Adaplation is shown to he one of the mechanisms which make the algae resistent. “Chlorophyll inactivation” is another. Experiments with the diatom Skeletonema costatum concerning adaptation to different temperatures have been performed. The fact that the alga has essentially the same rate of photosynthesis per cell at all light intensities at 20°C and 7°C, may be attributed to an increase of all the enzymes at the low temperature. The amount of protein per cell was twice as high at 7°C as at 20°C.     

Nitrogenase activity and dark CO2 fixation in the lichen Peltigera aphthosa Willd

The lichen Peltigera aphthosa consists of a fungus and green alga (Coccomyxa) in the main thallus and of a Nostoc located in superficial packets, intermixed with fungus, called cephalodia. Dark nitrogenase activity (acetylene reduction) of lichen discs (of alga, fungus and Nostoc) and of excised cephalodia was sustained at higher rates and for longer than was the dark nitrogenase activity of the isolated Nostoc growing exponentially. Dark nitrogenase activity of the symbiotic Nostoc was supported by the catabolism of polyglucose accumulated in the ligh and which in darkness served to supply ATP and reductant. The decrease in glucose content of the cephalodia paralleled the decline in dark nitrogenase activity in the presence of CO₂; in the absence of CO₂ dark nitrogenase activity declined faster although the rate of glucose loss was similar in the presence and absence of CO₂. Dark CO₂ fixation, which after 30 min in darkness represented 17 and 20% of the light rates of discs and cephalodia, respectively, also facilitated dark nitrogenase activity. The isolated Nostoc, the Coccomyxa and the excised fungus all fixed CO₂ in the dark; in the lichen most dark CO₂ fixation was probably due to the fungus. Kinetic studies using discs or cephalodia showed highest initial incorporation of ¹⁴CO₂ in the dark in to oxaloacetate, aspartate, malate and fumarate; incorporation in to alanine and citrulline was low; incorporation in to sugar phosphates, phosphoglyceric acid and sugar alcohols was not significant. Substantial activities of the enzymes phosphoenolpyruvate (PEP) carboxylase (EC 4.1.1.31) and carbamoyl-phosphate synthase (EC 2.7.2.5 and 2.7.2.9) were detected but the activities of PEP carboxykinase (EC 4.1.1.49) and PEP carboxyphosphotransferase (EC 4.1.1.38) were negligible. In the dark nitrogenase activity by the cephalodia, but not by the free-living Nostoc, declined more rapidly in the absence than in the presence of CO₂ in the gas phase. Exogenous NH ₄ ⁺ inhibited nitrogenase activity by cephalodia in the dark especially in the absence of CO₂ but had no effect in the light. The overall data suggest that in the lichen dark CO₂ fixation by the fungus may provide carbon skeletons which accept NH ₄ ⁺ released by the cyanobacterium and that in the absence of CO₂, NH ₄ ⁺ directly, or indirectly via a mechanism which involves glutamine synthetase, inhibits nitrogenase activity.Abbreviations CP carbamoyl phosphate - EDTA ethylenedi-amine tetraacetic acid - PEP phosphoenolpyruvate - RuBP ribulose 1,5 bisphosphate     

EFFECT OF TEMPERATURE ON THE SENSITIVITY OF NITROGENASE TO OXYGEN IN TWO HETEROCYSTOUS CYANOBACTERIA1

The effect of temperature and oxygen on nitrogenase activity in two heterocystous cyanobacteria, Anabaena variabilis Kütz. ATCC29413 and Nostoc sp. PCC7120, was investigated. The cyanobacteria were grown under a 12:12 light:dark (L:D) cycle at 27°C and were subsequently exposed to different temperatures (27, 36, 39, and 42°C) at different steady‐state O₂ concentrations (20, 10, 5, 0%). Light response curves of nitrogenase activity were recorded under each of these conditions using an online acetylene reduction assay combined with a sensitive laser photoacoustic ethylene detection method. The light response curves were fitted with the rectangular hyperbola model from which the model parameters N_m, N_d, and α were derived. In both strains, nitrogenase activity (N_tot = N_m + N_d) was the highest at 39°C–42°C and at 0% O₂. The ratio N_tot/N_d was 4.1 and 3.1 for Anabaena and Nostoc, respectively, indicating that respectively 25% and 33% of nitrogenase activity was supported by respiration (N_d). N_tot/N_d increased with decreasing O₂ concentration and with increasing temperature. Hence, each of these factors caused a relative increase in the light‐driven nitrogenase activity (N_m). These results demonstrate that photosynthesis and respiration both contribute to nitrogenase activity in Anabaena and Nostoc and that their individual contributions depend on both O₂ concentration and temperature as the latter may dynamically alter the flux of O₂ into the heterocyst.     

Characterization of a symbiotic,heterocystous, N2-fixing cyanobacterium fromCycas coralloid roots

A symbiotic, heterocystous, N₂-fixing blue-green alga, isolated from the coralloid roots of a xerophytic plant,Cycas revoluta, grew best in liquid medium supplemented with 4 mM NO ₃ ^– . Morphologically, the isolated alga was identical to that of the natural endophyte but the cell size had decreased markedly. The alga was heterotrophic. Intact coralloid roots had nearly 4 to 5 times more nitrogenase activity compared with natural- and laboratory-grown agla but nitrate reductase was inducible in both the forms. Plasmid(s) were found in both algal forms.     

Interactions between carbon dioxide and oxygen in the photosynthesis of three species of marine red macroalgae

Summary

Red algae have the highest known selectivity factor (S_rel) for CO₂ over O₂ of ribulose bisphosphate carboxylase-oxygenase (RUBISCO). This allows the prediction that a red alga relying on diffusive supply of CO₂ to RUBISCO from air-equilibrated solution should have less O₂ inhibition of photosynthesis than would an otherwise similar non-red alga with a lower S_rel of RUBISCO. Furthermore, RUBISCO shows an increased S_rel values at low temperatures. The prediction that O ₂inhibition of photosynthesis should be small for marine red algae relying on diffusive CO₂ entry growing in the North Sea with an annual temperature range of 4–16°C was tested in O₂ electrode experiments at 12°C. Phycodrys rubens and Plocamium cartilagineum, which rely on diffusive CO₂ entry showed, as predicted, only a small inhibition at lower inorganic C concentrations. Palmaria palmata, which has a CO₂ concentrating mechanism, had the expected negligible O ₂ inhibition of photosynthesis at any inorganic C concentration except (non-significantly) for saturating inorganic C.     

The Azolla-Anabaena azollae relationship

The heterocystous blue-green alga, Anabaena azollae, was isolated from the leaf cavities of the water fern, Azolla caroliniana, where it occurs as an endophyte. The isolated alga was capable of light dependent CO₂ fixation and acetylene reduction. Aerobic dark acetylene reduction occurred and was dependent upon endogenous substrates. Vegetative cells of the alga reduced nitro-blue tetrazolium chloride (NBT) to blue formazan. Heterocysts did not. Heterocysts reduced triphenyl tetrazolium chloride (TTC) to red formazan faster than vegetative cells. Reduction of TTC by both heterocysts and vegetative cells was much more rapid than has been reported for free-living heterocystous blue-green algae. Both NBT and TTC inhibited acetylene reduction and CO₂ fixation. The inhibition by TTC was more closely correlated to the time of exposure of the cells to the reagent and to the amount of deposition per cell than to the number of cells containing red formazan. No differential inhibition of acetylene reduction versus CO₂ fixation was observed. Autoradiography showed that CO₂ fixation occurred only in vegetative cells. Heterocysts caused a darkening of nuclear emulsions (chemography). This observation has been employed by others as an index of reducing activity in these cells. DCMU inhibited the acetylene reducing capacity of alga isolated from dark pretreated fronds more rapidly and to a greater extent than that in alga isolated from light pretreated fronds. Ammonia in excess of 5 mM was required before any inhibition of acetylene reduction was observed under either aerobic or anaerobic conditions in the light.     

Nitrogenase activity of the antarctic cyanobacteriumNostoc commune: Influence of temperature

Nitrogenase activity (C₂H₂ reduction) ofNostoc commune isolated from Schirmacher Oasis (Antarctica) was compared withNostoc muscorum, N. calcicola, Anabaena doliolum andGloeocapsa sp. The temperature profile of acetylene reduction (5–30 °C) forN. commune revealed (a) that the highest rate of nitrogenase activity was at 25±1 °C, (b) that it was low (69 %) in comparison withN. muscorum, and (c) that nitrogenase activity continued at lower temperatures, which was not evident for other cyanobacteria. The results suggest thatN. commune is adapted to lower temperatures in terms of nitrogen fixation.     

Effect of soil temperature on nitrogen fixation on roots of rice and reed

Summary The relation of nitrogenase activity (ethylene evolution) to soil temperature or incubation temperature of roots was determined on two genera of swamp plants, namely rice (Oryza sativa) cultivated in tropical climate and reed (Phragmites communis) grown in temperate regions. For both intact rice plants and excised rice roots the optimum temperature was 35°C. On excised roots nitrogenase activity responded more sensitivity to changes in temperature. In contrast to intact rice plants no ethylene evolution occurred on excised roots at 17 and 44°C. On reed roots temperature optimum was between 26 and 30°C which is clearly lower than on rice (35°C). The temperature range in which nitrogen fixation occurred was, however, similar to that of rice, although on a lower level. The results suggest a higher potential of the tropics for associative N₂ fixation, while in cooler climates the lower temperatures appear to be a major limiting factor.     

DIURNAL FLUCTUATION OF NITRATE REDUCTASE ACTIVITY IN THE MARINE RED ALGA GRACILARIA TENUISTIPITATA (RHODOPHYTA)1

The biochemical characteristics and diurnal changes in activity of the enzyme nitrate reductase (NR; EC 1.6.6.1) from the marine red alga Gracilaria tenuistipitata var. liui Zhang et Xia are described. Different assay conditions were tested to determine the stability of NR. The crude extract of G. tenuistipitata has a NR specific activity of 10.2 U.mg⁻¹, which is higher than the NR activities found for other algae, plants, and fungi. This NR is highly active at a slightly alkaline pH and is stable over a wide range of temperature, with an optimal activity at 20° C. The apical portions of the thallus contain 64.9 ± 6.6% of the total NR specific activity. The apparent Michaelis-Menten (K_m) constant found for KNO₃ was 197 μM, and it was 95 μM for NADH. The NR from G. tenuistipitata can be included in the NADH-specific group, because no activity was found when NADPH was used as an electron donor. In extracts of algae grown under either continuously dim light or a light-dark cycle, the activity of NR exhibits a daily rhythm, peaking at the middle of the light phase, when activity is 30-fold higher than during the night phase.     

DEPRESSION OF MASTIGOCLADUS LAMINOSUS (CYANOPHYTA) NITROGENASE ACTIVITY UNDER NORMAL SUNLIGHT INTENSITIES1

The effect of light intensity (PAR) on the nitrogenase activity of Mastigocladus laminosus Cohn was studied by the acetylene reduction technique. Benthic mat from a thermal stream, Hot River, in Yellowstone National Park was used in both experimental and in situ incubations. This hot spring maintained a mean pH of 7.0, was essentially isothermal (ca. 50°C), and had virtually no upstream to downstream physicochemical gradients (P > 0.05). Two surveys of the stream showed that nitrogenase of the M. laminosus mat was significantly more active (P > 0.02) under low light intensities than under high intensities, 252 and 712 μE · m^?2· s^?1, respectively. Maximum activity of Hot River Mastigocladus (268 nmol C₂H₄· mg Chl a^?1· h^?1) occurred at 50% full midday light intensities; the rates at low light (mean = 247 nmol C₂H₄· mg Chl a^?1· h^?1) were significantly (P > 0.001) greater than those at high light (mean = 106). The results indicate that M. laminosus nitrogenase activity is low light adapted and suggest that the temporal pattern for nitrogen fixation might be significantly different from that of thermophilic Calothrix.     

Relationship between carbohydrate movement and the symbiosis in lichens with green algae

Summary When isolated in pure culture, four genera of lichen algae were able to produce the polyol which is known to move from the alga to the fungus in lichens with these algae. This conclusion corrects earlier suggestions that the mobile polyol is only formed by the alga in the lichen thallus. Stichococcus produced sorbitol and it is therefore suggested that, in lichens with this alga, sorbitol moves between the symbionts. Hyalococcus and Stichococcus had a similar pattern of incorporation of H¹⁴CO ₃ ^- in the light, suggesting a close relationship between these algae which are only separated now on morphological grounds.The pattern of incorporation of H¹⁴CO ₃ ^- in the light into Cladonia cristatella and its alga (Trebouxia erici) in culture indicates that in the cultured algae more ¹⁴C was incorporated into ethanol insoluble substances and lipids and less into ribitol than in the lichen. The pattern in a joint culture of the alga and the fungus of C. cristatella was approximately intermediate between that of the lichen and the alga. However, only a small amount of ¹⁴C fixed by the alga reached the fungus in the joint culture, and it is therefore suggested that the presence of the fungus without morphological differentiation into a lichen thallus is not sufficient to promote the alga to release carbohydrate.     

Temperature responses of growth, photosynthesis, fatty acid and nitrate reductase in Antarctic and temperate Stichococcus

Stichococcus, a genus of green algae, distributes in ice-free areas throughout Antarctica. To understand adaptive strategies of Stichococcus to permanently cold environments, the physiological responses to temperature of two psychrotolerants, S. bacillaris NJ-10 and S. minutus NJ-17, isolated from rock surfaces in Antarctica were compared with that of one temperate S. bacillaris FACHB753. Two Antarctic Stichococcus strains grew at temperature from 4 to 25°C, while the temperate strain could grow above 30°C but could not survive at 4°C. The photosynthetic activity of FACHB753 at lower than 10°C was less than that of Antarctic algae. Nitrate reductase in NJ-10 and NJ-17 had its optimal temperature at 20°C, in comparison, the maximal activity of nitrate reductase in FACHB753 was found at 25°C. When cultured at 4–15°C a large portion of unsaturated fatty acids in the two Antarctic species was detected and the regulation of the degree of unsaturation of fatty acids by temperature was observed only above 15°C, though the content of the major unsaturated fatty acid αC18:3 in FACHB753 decreased with the temperatures elevated from 10 to 25°C. Elevated nitrate reductase activity and photosynthetic rates at low temperatures together with the high proportion of unsaturated fatty acids contribute to the ability of the Antarctic Stichococcus to thrive.     

Light Masking in the Field: An Experiment with Nocturnal and Diurnal Spiny Mice Under Semi-natural Field Conditions

Light masking has been studied almost exclusively in the laboratory. The authors populated four field enclosures with locally coexisting nocturnal Acomys cahirinus and diurnal A. russatus, and monitored their body temperatures (T_b) using implanted temperature-sensitive radio transmitters. A 3-h light pulse was initiated at the beginning of two consecutive nights; preceding nights were controls. A. cahirinus T_b and calculated activity levels decreased significantly during the light pulse, demonstrating a negative light masking response (light effect on T_b: ?0.32°C?±?0.15°C; average calculated activity records during the light pulse: 7?±?1.53, control: 9.8?±?1.62). Diurnal A. russatus did not respond to the light pulse. We conclude that light masking is not an artifact of laboratory conditions but represents a natural adaptive response in free-living populations. (Author correspondence: Shayroti@post.tau.ac.il)     

Dehydration rate and time of desiccation affect recovery of the lichenic algae <Emphasis Type="Italic">Trebouxia erici</Emphasis>: alternative and classical protective mechanisms

The mechanisms involved in desiccation tolerance of lichens and their photobionts are still poorly understood. To better understand these mechanisms we have studied dehydration rate and desiccation time in Trebouxia, the most abundant chlorophytic photobiont in lichen. Our findings indicate that the drying rate has a profound effect on the recovery of photosynthetic activity of algae after rehydration, greater than the effects of desiccation duration. The basal fluorescence (F′_o) values in desiccated algae were significantly higher after rapid dehydration, than after slow dehydration, suggesting higher levels of light energy dissipation in slow-dried algae. Higher values of PSII electron transport were recovered after rehydration of slow-dried Trebouxia erici compared to rapid-dried algae. The main component of non-photochemical quenching after slow dehydration was energy dependent (q _E), whereas after fast dehydration it was photoinhibition (q _I). Although q _E seems to play a role during desiccation recovery, no significant variations were detected in the xanthophyll cycle components. Desiccation did not affect PSI functionality. Classical antioxidant activities like superoxide dismutase or peroxidase decreased during desiccation and early recovery. Dehydrins were detected in the lichen-forming algae T. erici and were constitutively expressed. There is probably a minimal period required to develop strategies which will facilitate transition to the desiccated state in this algae. In this process, the xanthophyll cycle and classical antioxidant mechanisms play a very limited role, if any. However, our results indicate that there is an alternative mechanism of light energy dissipation during desiccation, where activation is dependent on a sufficiently slow dehydration rate.     

Ornithine cycle in Nostoc PCC 73102. Occurrence and localization of ornithine carbamoyl transferase, and the effects of external carbon and ornithine on nitrogenase activity and citrulline synthesis

Cells of free-living nitrogen-fixing Nostoc PCC 73102, a filamentous heterocystous cyanobacterium originally isolated from coralloid roots of the cycad Macrozamia. were examined for the presence of ornithine carbamoyl transferase (OCT) by native-PAGE/in situ activity stain, and SDS-PAGE/Western immunoblots. Transmission electron microscopy and immunocytological labeling were used to study the cellular and subcellular distribution of OCT in the Nostoc cells. Moreover, the effects of photoautotrophic and dark heterotrophic growth metabolism on growth, nitrogenase activity and in vivo citrulline synthesis were investigated. PAGE in combination with in situ activity staining demonstrated an in vitro active OCT with a molecular weight of approximately 80 kDa. SDS-PAGE/Western immunoblots revealed that a polypeptide with a molecular weight of approximately 38 kDa was immunologically related to OCT purified from pea (Pisum sativum L. cv. Alaska). Immunolocalization demonstrated that the OCT protein was located both in vegetative cells and heterocysts. Using the particle analysis of an image processor, the labeling associated with the photosynthetic vegetative cells was calculated to be 75.6 (± 5.5) gold particles μm^?2 compared with 62.0 (± 7.5) in the nitrogen-fixing heterocysts. Glucose and fructose stimulated both cyanobacterial growth and nitrogenase activity in light and darkness. Addition of exogenous ornithine decreased nitrogenase activity. In light grown cells, additions of glucose and fructose in combination with ornithine not only stimulated growth and nitrogenase activity but also in vivo citrulline synthesis, measured as ¹⁴CO₂-fixation into [¹⁴C]-citrulline. In darkness no stimulation was observed on in vivo citrulline synthesis. The substantial stimulation of nitrogenase activity by additions of external glucose and fructose, both in the light and in darkness, was not followed by a simultaneous stimulation of in vivo citrulline synthesis.     

Photosynthesis of Marine Macroalgae from Antarctica: Light and Temperature Requirements

The photosynthetic performance of macroalgae isolated in Antarctica was studied in the laboratory. Species investigated were the brown algae Himantothallus grandifolius, Desmarestia anceps, Ascoseira mirabilis, the red algae Palmaria decipiens, Iridaea cordata, Gigartina skottsbergii, and the green algae Enteromorpha bulbosa, Acrosiphonia arcta, Ulothrix subflaccida and U. implexa. Unialgal cultures of the brown and red algae were maintained at 0°C, the green algae were cultivated at 10°C. I_K values were between 18 and 53 μmol m^?2 s^?1 characteristic or low light adapted algae. Only the two Ulothrix species showed higher I_K values between 70 and 74 μmol m^?2 s^?1. Photosynthesis compensated dark respiration at very low photon fluence rates between 1.6 and 10.6 μmol m^?2 s^?1. Values of α were high: between 0.4 and 1.1 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the brown and red algae and between 2.1 and 4.9 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the green algal species. At 0°C P_max values of the brown and red algae ranged from 6.8 to 19.1 μmol O₂ g^?1 FW h^?1 and were similarly high or higher than those of comparable Arctic-cold temperate species. Optimum temperatures for photosynthesis were 5 to 10°C in A. mirabilis, 10°C in H. grandifolius, 15°C in G. skottsbergii and 20°C or higher in D. anceps and I. cordata. P: R ratios strongly decreased in most brown and red algae with increasing temperatures due to different Q₁₀ values for photosynthesis (1.4 to 2.5) and dark respiration (2.5 to 4.1). These features indicate considerable physiological adaptation to the prevailing low light conditions and temperatures of Antarctic waters. In this respect the lower depth distribution limits and the northern distribution boundaries of these species partly depend on the physiological properties described here.     

Components of natural gas resulting from thermal degradation of the blue-green alga (cyanobacterium)Oscillatoria tenuis

Cultures of the blue-green alga (cyanobacterium)Oscillatoria tenuis were used to simulate thermal degradation and gas formation by heating without oxygen at 250° and 350 °C for 100 h. Analysis through gas chromatography showed that the gases were mainly CH₄, C₂H₆, C₃H₈, iC₄ (isobutane), nC₄ (normal butane), iC₅ (isopentane), nC₅ (normal pentane), H₂, C0₂ and N₂. The volume of gases per g dry weight of alga was 44 ml at 250 °C and 100 ml at 350 °C. Alkane gas comprised only 2.04% of the total at 250 °C and rising to 40.0% at 350 °C. The fraction of C0₂ decreased from 83.3% at 250 °C to 40.0% at 350 °C. The quantity of alkane in the soluble organic matter doubled with rising temperature but the H/C atomic ratio in the ‘kerogen’, insoluble organic matter, decreased sharply. Infrared spectra of the ‘kerogen’ showed that the peak of adipose radical at 2900 cm⁻¹ disappeared gradually with rising temperature, which reflects the gradual break of CH₄ or C₂H₆ from ‘kerogen’. This demonstrates that insoluble organic matter rather than soluble organic matter in blue-green algae are the main sources of the gas alkanes in the process of simulated thermal degradation.