THE SHORT‐TERM EFFECT OF IRRADIANCE ON THE PHOTOSYNTHETIC PROPERTIES OF ANTARCTIC FAST‐ICE MICROALGAL COMMUNITIES1

Although sea‐ice represents a harsh physicochemical environment with steep gradients in temperature, light, and salinity, diverse microbial communities are present within the ice matrix. We describe here the photosynthetic responses of sea‐ice microalgae to varying irradiances. Rapid light curves (RLCs) were generated using pulse amplitude fluorometry and used to derive photosynthetic yield (Φ_PSII), photosynthetic efficiency (α), and the irradiance (E_k) at which relative electron transport rate (rETR) saturates. Surface brine algae from near the surface and bottom‐ice algae were exposed to a range of irradiances from 7 to 262 μmol photons · m^?2 · s^?1. In surface brine algae, Φ_PSII and α remained constant at all irradiances, and rETR_max peaked at 151 μmol photons · m^?2 · s^?1, indicating these algae are well acclimated to the irradiances to which they are normally exposed. In contrast, Φ_PSII, α, and rETR_max in bottom‐ice algae reduced when exposed to irradiances >26 μmol photons · m^?2 · s^?1, indicating a high degree of shade acclimation. In addition, the previous light history had no significant effect on the photosynthetic capacity of bottom‐ice algae whether cells were gradually exposed to target irradiances over a 12 h period or were exposed immediately (light shocked). These findings indicate that bottom‐ice algae are photoinhibited in a dose‐dependent manner, while surface brine algae tolerate higher irradiances. Our study shows that sea‐ice algae are able to adjust to changes in irradiance rapidly, and this ability to acclimate may facilitate survival and subsequent long‐term acclimation to the postmelt light regime of the Southern Ocean.     

EFFECT OF SEASONAL SEA ICE BREAKOUT ON THE PHOTOSYNTHESIS OF BENTHIC DIATOM MATS AT CASEY,ANTARCTICA1

Photosynthesis of marine benthic diatom mats was examined before and after sea ice breakout at a coastal site in eastern Antarctica (Casey). Before ice breakout the maximum under‐ice irradiance was between 2.5 and 8.2 μmol photons·m^?2·s^?1 and the benthic microalgal community was characterized by low E_k (12.1–32.3 μmol photons·m^?2·s^?1), low relETR_max (9.2–32.9), and high alpha (0.69–1.1). After breakout, 20 days later, the maximum irradiance had increased to between 293 and 840 μmol photons·m^?2·s^?1, E_k had increased by more than an order of magnitude (to 301–395 μmol photons·m^?2·s^?1), relETR_max had increased by more than five times (to 104–251), and alpha decreased by approximately 50% (to 0.42–0.68). During the same time interval the species composition of the mats changed, with a decline in the abundance of Trachyneis aspera (Karsten) Hustedt, Gyrosigma subsalsum Van Heurck, and Thalassiosira gracilis (Karsten) Hustedt and an increase in the abundance of Navicula glaciei Van Heurck. The benthic microalgal mats at Casey showed that species composition and photophysiology changed in response to the sudden natural increase in irradiance. This occurred through both succession shifts in the species composition of the mats and also an ability of individual cells to photoacclimate to the higher irradiances.     

THE INFLUENCE OF SALINITY AND TEMPERATURE COVARIATION ON THE PHOTOPHYSIOLOGICAL CHARACTERISTICS OF ANTARCTIC SEA ICE MICROALGAE1

The responses of sea ice microalgae to variation in ambient irradiance (0 to 150 μE · m^?2· s^?1), temperature (–6° to + 6° C), and salinity (0 to 100 ppt) were tested to determine whether these variables act independently or in concert to influence rates of microalgal photosynthesis. The photosynthetic efficiency and maximum photosynthetic rate for sea ice microalgae increased as a function of incubation temperature between -6° and + 6° C. Furthermore, photosynthetic efficiency, maximum photosynthetic rate, and quantum yield were greatest at salinities between SO and 50 ppt. In contrast, the mean specific absorption coefficients were lowest near seawater salinities, and the saturating irradiance, I_s, appeared to be inversely proportional to salinity. Results also suggest that the effects of salinity on the growth of sea ice microalgae are independent of those elicited by temperature or light, and that the functional relationship between salinity and light or temperature is multiplicative. This information is essential to the proper formulation of algorithms used to describe algal growth in environments where light, temperature, and salinity are changing simultaneously, such as within sea ice or within the water column at the marginal ice edge zone.     

PHOTOSYNTHESIS AND RESPIRATION OF THE CONCHOCELIS STAGE OF ALASKAN PORPHYRA (BANGIALES,RHODOPHYTA) SPECIES IN RESPONSE TO ENVIRONMENTAL VARIABLES1

Photosynthesis and respiration of three Alaskan Porphyra species, P. abbottiae V. Krishnam., P. pseudolinearis Ueda species complex (identified as P. “pseudolinearis” below), and P. torta V. Krishnam., were investigated under a range of environmental parameters. Photosynthesis versus irradiance (P–I) curves revealed that maximal photosynthesis (P_max), irradiance at maximal photosynthesis (I_max), and compensation irradiance (I_c) varied with salinity, temperature, and species. The P_max of Porphyra abbottiae conchocelis varied between 83 and 240 μmol O₂ · g dwt^?1 · h^?1 (where dwt indicates dry weight) at 30–140 μmol photons · m^?2 · s^?1 (I_max) depending on temperature. Higher irradiances resulted in photoinhibition. Maximal photosynthesis of the conchocelis of P. abbottiae occurred at 11°C, 60 μmol photons · m^?2·s^?1, and 30 psu (practical salinity units). The conchocelis of P. “pseudolinearis” and P. torta had similar P_max values but higher I_max values than those of P. abbottiae. The P_max of P. “pseudolinearis” conchocelis was 200–240 μmol O₂ · g dwt^?1 · h^?1 and for P. torta was 90–240 μmol O₂ · g dwt^?1 · h^?1. Maximal photosynthesis for P. “pseudolinearis” occurred at 7°C and 250 μmol photons · m^?2 · s^?1 at 30 psu, but P_max did not change much with temperature. Maximal photosynthesis for P. torta occurred at 15°C, 200 μmol photons · m^?2 · s^?1, and 30 psu. Photosynthesis rates for all species declined at salinities <25 or >35 psu. Estimated compensation irradiances (I_c) were relatively low (3–5 μmol · photons · m^?2 · s^?1) for intertidal macrophytes. Porphyra conchocelis had lower respiration rates at 7°C than at 11°C or 15°C. All three species exhibited minimal respiration rates at salinities between 25 and 35 psu.     

COMPARISON OF DEVELOPMENT AND PHOTOSYNTHETIC GROWTH FOR FILAMENT CLUMPS AND REGENERATED MICROPLANTLET CULTURES OF AGARDHIELLA SUBULATA (RHODOPHYTA, GIGARTINALES)

Two axenic, in vitro liquid suspension cultures were established for Agardhiella subulata (C. Agardh) Kraft et Wynne, and their growth characteristics were compared. This study illustrated how reliable routes for the development of suspension cultures of macrophytic red algae of terete thallus morphology can be achieved for biotechnology applications. Undifferentiated filament clumps of 2–8 mm diameter were established by induction of callus-like tissue from thallus explants, and lightly branched microplantlets of 2–10 mm length were established by regeneration of filament clumps. The filament clumps were susceptible to regeneration. Adventitious shoot formation was reliably induced from 40% to 70% of the filament clumps by gentle mixing at 100 rev min^?1 on an orbital shaker. The specific growth rate of the microplantlets was higher than the filament clumps in nonagitated well plate culture (4%–6% per day for microplantlets vs. 2%–3% per day for filament clumps) at 24° C and 8–36 μmol photons·m^?2·s^?1 irradiance (10:14 h LD cycle) when grown on ASP12 artificial seawater medium at pH 8.6–8.9 with 20%–25% per day medium replacement. Oxygen evolution rate vs. irradiance measurements showed that relative to the filament clumps, microplantlets had a higher maximum specific oxygen evolution rate (P_o,max= 0.181 ± 0.035 vs. 0.130 ± 0.023 mmol O₂·g^?1 dry cell mass·h^?1), but comparable respiration rate (Q_o= 0.040 ± 0.013 vs. 0.033 ± 0.017 mmol O₂·g^?1 dry cell mass·h^?1), compensation point (I_c= 3.8 ± 2.4 vs. 5.7 ± 1.2 μmol photons·m^?2·s^?1), and light intensity at 63.2% of saturation (I_k= 17.5 ± 3.9 vs. 14.9 ± 2.6 μmol photons·m^?2·s^?1). The microplantlet culture was more suitable for suspension culture development than the filament clump culture because it was morphologically stable and exhibited higher growth rates.     

PHOTOSYNTHESIS-IRRADIANCE RELATIONSHIPS IN SEA ICE MICROALGAE FROM McMURDO SOUND,ANTARCTICA1

Sea ice microalgae in McMurdo Sound, Antarctica were examined for photosynthesis-irradiance relationships and for the extent and time course of their photoadaptation to a reduction in in situ irradiance. Algae were collected from the bottom centimeter of coarse-grained congelation ice in an area free of natural snow cover. Photosynthetic rate was determined in short term (1 h) incubations at ?2° C over a range of irradiance from 0 to 286 μE·m^?2·s^?1. Assimilation numbers were consistently below 0.1 mg C·mg chl a^?1·h^?1. The I_k's³ averaged only 7 μE·m^?2·s^?1, and photosynthesis was inhibited at irradiances above 25 μE·m^?2·s^?1. Photosynthetic parameters of the ice algal community were examined over a nine day period following the addition of 4 cm of surface snow while a control area remained snow-free. A reduction of 40% in P_m^B relative to the control occurred after two days of snow cover; α, β, I_k, and I_m were not significantly altered. Low assimilation numbers and constant standing crop size, however, suggested that the algal bloom may have already reached stationary growth phase, possibly minimizing their photoadaptive response.     

PHOTOSYNTHESIS AND COLD ACCLIMATION: MOLECULAR EVIDENCE FROM A POLAR DIATOM1

The psychrophilic diatom Fragilariopsis cylindrus (Grunow) Krieger in Helmcke & Krieger was used to investigate photosynthesis and growth under freezing temperatures. Gene expression during a temperature shift from +5° C to ?1.8° C was studied under 3 and 35 μmol photons·m^?2·s^?1 by using a macroarray. These measurements were paralleled by determination of fluorescence induction at PSII and pigment analysis. The shift to ?1.8° C at 35 μmol photons·m^?2·s^?1 caused a marginal decrease of photosynthetic quantum yield (F_v/F_m) from 0.61 to 0.52 with fast recovery after 1 day. The ratio of chl c to chl a increased from 3.1 to 5.5, and the ratio of diatoxanthin to diadinoxanthin increased from 0.7 to 5.0. Genes encoding proteins of PSII (psbA, psbC) and for carbon fixation (rbcL) were down‐regulated, whereas genes encoding chaperons (hsp70) and genes for plastid protein synthesis and turnover (elongation factor EfTs, ribosomal protein rpS4, ftsH protease) were up‐regulated. In contrast, cold exposure at 3 μmol photons·m^?2·s^?1 induced a marginal increase in F_v/F_m from 0.61 to 0.63 and a strong increase in fucoxanthin concentrations from 0.04 up to 0.12 pg·cell^?1. This was paralleled by up‐regulation of fcp genes. The ratio of chl c to chl a also increased from 3.1 to 4.2, as did the ratio of diatoxanthin to diadinoxanthin from 0.7 to 2.2. Down‐regulation of psbA, psbC, and rbcL could also be measured but not up‐regulation of hsp70, EfTs, rpS4, and the ftsH protease. The latter genes are probably necessary to avoid cold shock photoinhibition only at higher light intensities.     

EFFECTS OF LIGHT AND TEMPERATURE ON GROWTH,NITRATE UPTAKE,AND TOXIN PRODUCTION OF TWO TROPICAL DINOFLAGELLATES: ALEXANDRIUM TAMIYAVANICHII AND ALEXANDRIUM MINUTUM (DINOPHYCEAE)1

The two tropical estuarine dinoflagellates, Alexandrium tamiyavanichii Balech and A. minutum Halim, were used to determine the ecophysiological adaptations in relation to their temperate counterparts. These species are the two main causative organisms responsible for the incidence of paralytic shellfish poisoning (PSP) in Southeast Asia. The effects of light (10, 40, 60, and 100 μmol photons·m^?2·s^?1) and temperature (15, 20, and 25°C) on the growth, nitrate assimilation, and PST production of these species were investigated in clonal batch cultures over the growth cycle. The growth rates of A. tamiyavanichii and A. minutum increased with increasing temperature and irradiance. The growth of A. tamiyavanichii was depressed at lower temperature (20°C) and irradiance (40 μmol photons·m^?2·s^?1). Both species showed no net growth at 10 μmol photons·m^?2·s^?1 and a temperature of 15°C, although cells remained alive. Cellular toxin quotas (Q_t) of A. tamiyavanichii and A. minutum varied in the range of 60–180 and 10–42 fmol PST·cell^?1, respectively. Toxin production rate, R_tox, increased with elevated light at both 20 and 25°C, with a pronounced effect observed at exponential phase in both species (A. tamiyavanichii, r²=0.95; A. minutum, r²=0.96). Toxin production rate also increased significantly with elevated temperature (P<0.05) for both species examined. We suggest that the ecotypic variations in growth adaptations and toxin production of these Malaysian strains may reveal a unique physiological adaptation of tropical Alexandrium species.     

CULTURE OF THE TERRESTRIAL CYANOBACTERIUM,NOSTOC FLAGELLIFORME (CYANOPHYCEAE), UNDER AQUATIC CONDITIONS1

Both colonies and free‐living cells of the terrestrial cyanobacterium, Nostoc flagelliforme (Berk. & Curtis) Bornet & Flahault, were cultured under aquatic conditions to develop the techniques for the cultivation and restoration of this endangered resource. The colonial filaments disintegrated with their sheaths ruptured in about 2 days without any desiccating treatments. Periodic desiccation played an important role in preventing the alga from decomposing, with greater delays to sheath rupture with a higher frequency of exposure to air. The bacterial numbers in the culture treated with seven periods of desiccation per day were about 50% less compared with the cultures without the desiccation treatment. When bacteria in the culture were controlled, the colonial filaments did not disintegrate and maintained the integrity of their sheath for about 20 days even without the desiccation treatments, indicating the importance of desiccation for N. flagelliforme to prevent them from being disintegrated by bacteria. On the other hand, when free‐living cells obtained from crushed colonial filaments were cultured in liquid medium, they developed into single filaments with sheaths, within which multiple filaments were formed later on as a colony. Such colonial filaments were developed at 15, 25, and 30° C at either 20 or 60 μmol photons·m^?2·s^?1; colonies did not develop at 180 μmol photons·m^?2·s^?1, though this light level resulted in the most rapid growth of the cells. Conditions of 60 μmol photons·m^?2·s^?1 and 25° C appeared to result in the best colonial development and faster growth of the sheath‐held colonies of N. flagelliforme when cultured indoor under aquatic conditions.     

PRIMARY PRODUCTION OF CRUSTOSE CORALLINE RED ALGAE IN A HIGH ARCTIC FJORD1

Crustose coralline algae occupied ～1%–2% (occasionally up to 7%) of the sea floor within their depth range of 15–50 m, and they were the dominant encrusting organisms and macroalgae beyond 20 m depth in Young Sound, NE Greenland. In the laboratory, oxygen microelectrodes were used to measure net photosynthesis (P) versus downwelling irradiance (E_d) and season for the two dominant corallines [Phymatolithon foecundum (Kjellman) Düwel et Wegeberg 1996 and Phymatolithon tenue (Rosenvinge) Düwel et Wegeberg 1996] representing> 90% of coralline cover. Differences in P‐E_d curves between the two species, the ice‐covered and open‐water seasons, or between specimens from 17 and 36 m depth were insignificant. The corallines were low light adapted, with compensation irradiances (E_c) averaging 0.7–1.8 μmol photons·m ^{? 2}·s ^{? 1} and light adaptation (E_k) indices averaging 7–17 μmol photons·m ^{? 2}·s ^{? 1}. Slight photoinhibition was evident in most plants at irradiances up to 160 μmol photons·m ^{? 2}·s ^{? 1}. Photosynthetic capacity (P_m) was low, averaging 43–67 mmol O₂·m ^{? 2} thallus·d ^{? 1} (～250–400 g C·m ^{? 2} thallus·yr ^{? 1}). Dark respiration rates averaged ～5 mmol O₂·m ^{? 2} thallus·d ^{? 1}. In ice covered periods, E_d at 20 m depth averaged ～1 μmol photons·m ^{? 2}·s ^{? 1}, with daily maxima of 2–3 μmol photons·m ^{? 2}·s ^{? 1}. During the open water season, E_d at 20 m depth averaged ～7 μmol photons·m ^{? 2}·s ^{? 1} with daily maxima of ～30 μmol photons·m ^{? 2}·s ^{? 1}. Significant net primary production of corallines was apparently limited to the 2–3 months with open water, and the small contribution of corallines to primary production seems due to low P_m values, low in situ irradiance, and their relatively low abundance in Young Sound.     

HETEROGENEITY OF OXYGEN PRODUCTION AND CONSUMPTION IN A PHOTOSYNTHETIC MICROBIAL MAT AS STUDIED BY PLANAR OPTODES

By applying planar optodes and imaging techniques to a benthic photosynthetic mat, we demonstrated an extensive vertical and horizontal variation in O₂ concentrations, O₂ consumption, and O₂ production. In light, the oxic zone could be divided into three horizons: 1) an upper zone dominated by diatoms that had a moderate net O₂ production, 2) another zone dominated by Microcoleus-like cyanobacteria with a high net O₂ production, and 3) a lower zone with disintegrating microalgae and cyanobacteria with a high O₂ consumption rate. From the O₂ images, the net O₂ production/consumption was calculated at a spatial resolution of 130 μM. This allowed us to identify microsites with high rates of O₂ turnover within the photic zone. Sites with high net O₂ consumption (>1.5 nmol·cm^?3·s^?1) were typically situated next to sites with a relatively high net production (>2 nmol·cm^?3·s^?1), revealing a mosaic in which the highest O₂ consumption sites were surrounded by the highest O₂ production sites. This suggested a tight spatial coupling between production and consumption of O₂ within the photic zone. Light stimulated the O₂ consumption within the photic zone. At irradiances above 400 μmol photons·m^?2·s^?1, the stimulated O₂ production was almost completely balanced by enhanced O₂ consumption at microsites exhibiting net consumption of O₂ even at maximum irradiance (578 μmol photons·m^?2·s^?1). Our observations strongly supported the idea that light-stimulated respiration was caused by stimulated heterotrophic activity fueled by organic carbon leakage from the phototrophs. Despite microsites with high net O₂ consumption, anoxic microniches were not encountered in the investigated mat. Images of gross photosynthetic rates also revealed an extensive horizontal variation in gross rates, with microsites of low or no photosynthesis within the otherwise photic zone. Calculations based on the obtained images revealed that at maximum light (578 μmol photons·m^?2·s^?1), 90% of the O₂ produced was consumed within the photic zone. The presented data demonstrate the great potential offered by planar optode for studies of benthic photosynthetic communities.     

The effect of irradiance and temperature responses and the phenology of a native alga,Undaria pinnatifida (Laminariales), at the southern limit of its natural distribution in Japan

Phenology, irradiance, and temperature characteristics of an edible brown alga, Undaria pinnatifida (Laminariales), were examined from the southernmost natural population in Japan, both by culturing gametophytes and examining the photosynthetic activity of sporophytes using dissolved oxygen sensors and pulse amplitude-modulated chlorophyll fluorometer (IMAGING-PAM). Our surveys confirmed that sporophytes were present between winter and early summer, but absent by July. IMAGING-PAM experiments were used to measure maximum effective quantum yield (ΦII at 0 μmol photons m^?2 s^?1) for each of 14 temperatures (8–36 °C). Oxygen production was also determined over a coarser temperature gradient. Net photosynthesis and ΦII (at 0 μmol photons m^?2 s^?1) were observed to be temperature-dependent; the maximum ΦII was estimated to be 0.67, occurred at 21.2 °C, and was nearly identical to the optimal temperature of the net photosynthetic rate (21.7 °C). A net photosynthesis–irradiance (P–E) model revealed that saturation irradiance (E _k) was 119.5 μmol photons m^?1 s^?1, and the compensation irradiance (E _c) was 17.4 μmol photons m^?1 s^?1. Culture experiments on the gametophytes revealed that most individuals could not survive temperatures over 28 °C and that growth rates were severely inhibited. Based on our observations, temperatures greater than 20 °C are likely to influence photosynthetic activity and gametophyte survival, and therefore, it is possible that this species might become locally extinct if seawater temperatures in this region continue to rise.     

SHADE ADAPTED BENTHIC DIATOMS BENEATH ANTARCTIC SEA ICE1

A dense community of shade adapted microalgae dominated by the diatom Trachyneis aspera is associated with a siliceous sponge spicule mat in McMurdo Sound, Antarctica. Diatoms at a depth of 20 to 30 m were found attached to spicule surfaces and in the interstitial water between spicules. Ambient irradiance was less than 0.6 μE · m^?2· s^?1 due to light attenuation by surface snow, sea ice, ice algae, and the water column. Photosynthesis-irradiance relationships determined by the uptake of NaH¹⁴CO₃ revealed that benthic diatoms beneath annual sea ice were light-saturated at only 11 μE·m^?2·s^?1, putting them among the most shade adapted microalgae reported. Unlike most shade adapted microalgae, however, they were not photoinhibited even at irradiances of 300 μE·m^?2·s^?1. Although in situ primary production by benthic diatoms was low, it may provide a source of fixed carbon to the abundant benthic invertebrates when phytoplankton or ice algal carbon is unavailable.     

PHOTOINHIBITION OF MECHANICALLY STIMULABLE BIOLUMINESCENCE IN THE AUTOTROPHIC DINOFLAGELLATE CERATIUM FUSUS (PYRROPHYTA)1

Ceratium fusus (Ehrenb.) Dujardin was exposed to light of different wavelengths and photon flux densities (PFDs) to examine their effects on mechanically stimulable bioluminescence (MSL). Photoinhibition of MSL was proportional to the logarithm of PFD. Exposure to I μmol photons·m^?2s^?1 of broadband blue light (ca. 400–500 nm) produced near-complete photoinhibition (≥90% reduction in MSL) with a threshold at ca. 0.01 μmol photons·m^?2·s^?1. The threshold of photoinhibition was ca. an order of magnitude greater for both broadband green (ca. 500–580 nm) and red light (ca. 660–700 nm). Exposure to narrow spectral bands (ca. 10 nm half bandwidth) from 400 and 700 nm at a PFD of 0.1 μmol photons·m^?2·s^?1 produced a maximal response of photoinhibition in the blue wavelengths (peak ca. 490 nm). A photoinhibition response (≥ 10%) in the green (ca. 500–540 nm) and red wavelengths (ca. 680 nm) occurred only at higher PFDs (1 and 10 μmol photons·m^?2·s^?1). The spectral response is similar to that reported for Gonyaulax polyedra Stein and Pyrocystis lunula Schütt and unlike that of Alexandrium tamarense (Lebour) Balech et Tangen. The dinoflagellate's own bioluminescence is two orders of magnitude too low to result in self-photoinhibition. The quantitative relationships developed in the laboratory predict photoinhibition of bioluminescence in populations of C. fusus in the North Atlantic Ocean.     

PHOTOSYNTHESIS-IRRADIANCE RELATIONSHIPS IN PHYTOPLANKTON FROM THE PHYSICALLY STABLE WATER COLUMN OF A PERENNIALLY ICE-COVERED LAKE (LAKE BONNEY,ANTARCTICA)1

The perennially ice-covered lakes of Antarctica have hydrodynamically stable water columns with a number of vertically distinct phytoplankton populations. We examined the photosynthesis-irradiance characteristics of phytoplankton from four depths of Lake Bonney to determine their physiological condition relative to vertical gradients in irradiance and temperature. All populations studied showed evidence of extreme shade adaptation, including low I_k values (15–45 μE · m^?2· s^?1) and extremely low maximal photosynthetic rates (P^B_m less than 0.3 μg C ·μg chl a^?1· h^?1). Photosynthetic rates were controlled by temperature as well as light variations with depth. Lake Bonney has an inverted temperature profile within the trophogenic zone that increased from 0° C at the ice-water interface to 6° C from 10 to 18 m. Deeper phytoplankton (10 m and 17 m) were found to have photosynthetic capacities (P^B_m) and efficiences (α) three to five times higher than those at the ice-water interface. However, Q₁₀ values were only ca. 2 for P^B_m (no temperature dependence was evident for α), suggesting that a simple temperature response cannot explain all the differences between populations. Lake Bonney phytoplankton (primarily cryptophytes and chlorophytes) had photosynthetic characteristics similar to diatoms from other physically stable environments (e.g. sea ice, benthos) and may be ecologically analogous to multiple deep chlorophyll maxima.     

EFFECTS OF LIGHT AND TEMPERATURE ON NET PHOTOSYNTHESIS AND DARK RESPIRATION OF GAMETOPHYTES AND EMBRYONIC SPOROPHYTES OF MACROCYSTIS PYRIFERA1

The rates of net photosynthesis as a function of irradiance and temperature were determined for gametophytes and embryonic sporophytes of the kelp, Macrocystis pyrifera (L.) C. Ag. Gametophytes exhibited higher net photosynthetic rates based on oxygen and pH measurements than their derived embryonic sporophytes, but reached light saturation at comparable irradiance levels. The net photosynthesis of gametophytes reached a maximum of 66.4 mg O₂ g dry wt^?1 h^?1 (86.5 mg CO₂ g dry wt^?1 h^?1), a value approximately seven times the rate reported previously for the adult sporophyte blades. Gametophytes were light saturated at 70 μE m^?2 s^?1 and exhibited a significant decline in photosynthetic performance at irradiances 140 μE m^?1 s^?1. Embryonic sporophytes revealed a maximum photosynthetic capacity of 20.6 mg O₂ g dry wt^?1 h^?1 (25.3 mg CO₂ g dry wt^?1 h^?1), a rate about twice that reported for adult sporophyte blades. Embryonic sporophytes also became light saturated at 70 μE m^?2 s^?1, but unlike their parental gametophytes, failed to exhibit lesser photosynthetic rates at the highest irradiance levels studied; light compensation occurred at 2.8 μE m^?2 s^?1. Light-saturated net photosynthetic rates of gametophytes and embryonic sporophytes varied significantly with temperature. Gametophytes exhibited maximal photosynthesis at 15° to 20° C, whereas embryonic sporophytes maintained comparable rates between 10° and 20° C. Both gametophytes and embryonic sporophytes declined in photosynthetic capacity at 30° C. Dark respiration of gametophytes was uniform from 10° to 25° C, but increased six-fold at 30° C; the rates for embryonic sporophytes were comparable over the entire range of temperatures examined. The broader light and temperature tolerances of the embryonic sporophytes suggest that this stage in the life history of M. pyrifera is well suited for the subtidal benthic environment and for the conditions in the upper levels of the water column.     

THE INFLUENCE OF FLUCTUATING LIGHT ON DIVERSITY AND SPECIES NUMBER OF NUTRIENT‐LIMITED PHYTOPLANKTON1

The influence of fluctuating light on diversity and species number of a natural phytoplankton assemblage competing for nutrients was investigated for 48 days under semicontinuous culture conditions. Light conditions were either changed periodically from high (65 μmol photons·m^?2·s^?1) to low intensity (15 μmol photons·m^?2·s^?1) at intervals of 1, 3, 6, and 12 days or fixed at constant light conditions of intermediate intensity (40 μmol photons·m^?2·s^?1). Fluctuating light at intervals of 1–12 days significantly affected phytoplankton diversity. The development of phytoplankton communities differed in treatments with different light regimes. In treatments with long light intervals, species abundance oscillated with the light phases. Differences in the temporal development of phytoplankton communities resulted in hump‐shaped relations between the interval length of the light phases and both species number and diversity index and can be explained by the intermediate disturbance hypothesis. Fluctuating light tends to sustain phytoplankton diversity under nutrient limitation if the light regime changes in the order of several days. This indicates that temporal changes in weather regime are important in preventing competitive exclusion of phytoplankton species in nature.     

PRODUCTIVITY OF THE FILAMENTOUS ALGA PITHOPHORA OEDOGONIA (CHLOROPHYTA) IN SURREY LAKE,INDIANA1

Biomass, akinete numbers, net photosynthesis, and respiration of Pithophora oedogonia were monitored over two growing seasons in shallow Surrey Lake, Indiana. Low rates of photosynthesis occurred from late fall to early spring and increased to maximum levels in late spring to summer (29–39 mgO₂·g^?1 dry wt·h^?1). Areal biomass increased following the rise in photosynthesis and peaked in autumn (163–206g dry wt·m^?2). Photosynthetic rates were directly correlated with temperature, nitrogen, and phosphorus over the entire annual cycle and during the growing season. Differences in photosynthetic activity and biomass between the two growing seasons (1980 and 1981) were apparently related to higher, early spring temperatures and higher levels of NO₃-N and PO₄-P in 1981. Laboratory investigations of temperature and light effects on Pithophora photosynthesis and respiration indicated that these processes were severely inhibited below 15°C. The highest P_max value occurred at 35°C (0.602 μmol O₂·mg^?1 chl a·min^?1). Rates of dark respiration did not increase above 25°C thus contributing to a favorable balance of photosynthetic production to respiratory utilization at high temperatures. Light was most efficiently utilized at 15°C as indicated by minimum values of I_k(47 μE·m^?2·s^?1) and I_c (6 μE·m^?2·s^?1). Comparison of P. oedogonia and Cladophora glomerata indicated that the former was more tolerant of temperatures above 30°C. Pithophora's tolerance of high temperature and efficient use of low light intensity appear to be adaptive to conditions found within the dense, floating algal mats and the shallow littoral areas inhabited by this filamentous alga.     

Effect of irradiance and temperature on the photosynthesis of a cultivated red alga,Pyropia tenera (= Porphyra tenera), at the southern limit of distribution in Japan

The effect of irradiance and temperature on the photosynthesis of the red alga, Pyropia tenera, was determined for maricultured gametophytes and sporophytes collected from a region that is known as one of the southern limits of its distribution in Japan. Macroscopic gametophytes were examined using both pulse‐amplitude modulated fluorometry and/or dissolved oxygen sensors. A model of the net photosynthesis–irradiance (P‐E) relationship of the gametophytes at 12°C revealed that the net photosynthetic rate quickly increased at irradiances below the estimated saturation irradiance of 46 μmol photons m^?2 s^?1, and the compensation irradiance was 9 μmol photons m^?2 s^?1. Gross photosynthesis and dark respiration for the gametophytes were also determined over a range of temperatures (8–34°C), revealing that the gross photosynthetic rates of 46.3 μmol O₂ mg_chl‐a^?1 min^?1 was highest at 9.3 (95% Bayesian credible interval (BCI): 2.3–14.5)°C, and the dark respiration rate increased at a rate of 0.93 μmol O₂ mg_chl‐a^?1 min^?1°C^?1. The measured dark respiration rates ranged from ?0.06 μmol O₂ mg_chl‐a^?1 min^?1 at 6°C to ?25.2 μmol O₂ mg_chl‐a^?1 min^?1 at 34°C. The highest value of the maximum quantum yield (Fv/Fm) for the gametophytes occurred at 22.4 (BCI: 21.5–23.3) °C and was 0.48 (BCI: 0.475–0.486), although those of the sporophyte occurred at 12.9 (BCI: 7.4–15.1) °C and was 0.52 (BCI: 0.506–0.544). This species may be considered well‐adapted to the current range of seawater temperatures in this region. However, since the gametophytes have such a low temperature requirement, they are most likely close to their tolerable temperatures in the natural environment.     

LIGHT AND TEMPERATURE AS FACTORS REGULATING SEASONAL GROWTH AND DISTRIBUTION OF ULOTHRIX ZONATA (ULVOPHYCEAE)1

Ulothrix zonata (Weber and Mohr) Kütz. is an unbranched filamentous green alga found in rocky littoral areas of many northern lakes. Field observations of its seasonal and spatial distribution indicated that it should have a low temperature and a high irradiance optimum for net photosynthesis, and at temperatures above 10°C it should show an increasingly unfavorable energy balance. Measurements of net photosynthesis and respiration were made at 56 combinations of light and temperature. Optimum conditions were 5°C and 1100 μE·m^?2·s^?1 at which net photosynthesis was 16.8 mg O₂·g^?1·h^?1. As temperature increased above 5° C optimum irradiance decreased to 125 μE·m^?2·s^?1 at 30°C. Respiration rates increased with both temperature and prior irradiance. Light-enhanced respiration rates were significantly greater than dark respiration rates following irradiance exposures of 125 μE·m^?2·s^?1 or greater. Polynomials were fitted to the data to generate response surfaces. Polynomial equations represent statistical models which can accurately predict photosynthesis and respiration for inclusion in ecosystem models.