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.     

SHORT‐TERM EFFECT OF TEMPERATURE ON THE PHOTOKINETICS OF MICROALGAE FROM THE SURFACE LAYERS OF ANTARCTIC PACK ICE1

Microalgae growing within brine channels (85 psu salinity) of the surface ice layers of Antarctic pack ice showed considerable photosynthetic tolerance to the extreme environmental condition. Brine microalgae exposed to temperatures above ?5°C and at irradiances up to 350 μmol photons·m^?2·s^?1 showed no photosynthetic damage or limitations. Photosynthesis was limited (but not photoinhibited) when brine microalgae were exposed to ?10°C, provided the irradiance remained under 50 μmol photons·m^?2·s^?1. The highest level of photosynthetic activity (maximum relative electron transport rate [rETR_max]) in brine microalgae growing within the surface layer of sea ice was at approximately 18 μmol electrons·m^?2·s^?1, which occurred at ?1.8°C. Effective quantum yield of PSII and rETR_max of the halotolerant brine microalgae exhibited a temperature‐dependent pattern, where both parameters were higher at ?1.8°C and lower at ?10°C. Relative ETR_max at temperatures above ?5°C were stable across a wide range of irradiance.     

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.     

CRITICAL IRRADIANCE LEVELS AND THE INTERACTIVE EFFECTS OF QUANTUM IRRADIANCE AND DOSE ON GAMETOGENESIS IN THE GIANT KELP,MACROCYSTIS PYRIFERA1

Gametophytes of Macrocystis pyrifera (L.) C. Ag. were cultured under a series of quantum irradiances in three photoperiod regimes. The quantum irradiances in each photoperiod were adjusted to provide equal daily irradiation dosages between photoperiods which allowed a critical examination of the interactions between quantum irradiance and quantum dose in determining gametophyte fertility. The lowest quantum irradiance which stimulated gametogenesis in more than 50% of the female gametophytes was 5 μE·m^?2·s^?1. The saturating irradiance was ca. 10 μE·m^?2·s^?1 at photoperiods of 12 h or greater. In terms of daily quantum dose, the lowest dose at which greater than 50% gametogenesis occurred was 0.2 E·m^?2·d^?1. However, this critical quantum dose was higher (0.4 E·m^?2·d^?1) when instantaneous irradiances were less than 5 μE·m^?2·s^?1. The saturation quantum dose was also affected by the rate at which the quantum dose was received and varied from 0.4 to 0.8 E·m^?2·d^?1. Gametophytes in all three photoperiods reached 100% fertility at quantum irradiances above 5 μE·m^?2·s^?1. Photoperiod effects were small and could be accounted for by quantum dosage effects.     

PHYSIOLOGICAL RESPONSES OF ANABAENA VARIABILIS (CYANOPHYCEAE) TO INSTANTANEOUS EXPOSURE TO VARIOUS COMBINATIONS OF LIGHT INTENSITY AND TEMPERATURE1

Light intensity and temperature interactions have a complex effect on the physiological process rates of the filamentous bluegreen alga Anabaena variabilis Kütz. The optimum temperature for photosynthesis increased with increasing light intensity from 10°C at 42 μE·m^?2·s^?1 to 35°C at 562 μE·m^?2·s^?1. The light saturation parameter, I_K, increased with increasing temperatures. The maximum photosynthetic rate (2.0 g C·g dry wt.^?1·d^?1) occurred at 35°C and 564 μE·m^?2·s^?1. At 15°C, the maximum rate was 1.25 g C·g dry wt.^?1·d^?1 at 332 μE·m^?2·s^?1. The dark respiration rate increased exponentially with temperature. Under favorable conditions of light intensity and temperature the percent of extracellular release of dissolved organic carbon was less than 5% of the total C fixed. This release increased to nearly 40% under combinations of low light intensity and high temperature. A mathematical model was developed to simulate the interaction of light intensity and temperature on photosynthetic rate. The interactive effects were represented by making the light-saturation parameters a function of temperature.     

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.     

THF INFLUENCE OF IRRADIANCE ON THF BIOCHEMICAL COMPOSITION OF THE PRYMNESIOPHYTE ISOCHRYSIS SP. (CLONE T-ISO)1

The effects of irradiance on the biochemical composition of the prymnesiophyte microalga, Isochrysis sp. (Parke; clone T-ISO), a popular species for mariculture, were examined. Cultures were grown under a 12:12 h light: dark (L:D) regime at five irradiances ranging from 50 to 1000 μE·m ²·s^?1 and harvested at late-logarithmic phase for analysis of biochemical composition. Gross composition varied aver the range of irradiances. The highest levels of protein were present in cells from cultures grown at 100 and 250 μE·m ³·s¹, and minimum levels of carbohydrate and lipid occurred at 50 μE·m^?2·s^?1. Because the cell dry weight was reduced at lower irradiances, different trends were evident when results were expressed as percentage of dry weights. Protein percentages were highest at Wand 100 μE·m^?2·s^?1 and carbohydrate at 100 μE·m^?2·s^?1. The composition of amino acids did not differ over the range of irradiances. Glutamate and aspartate were always present in high proportions (9.0–13.5%); histidine. methionine, tryptophan, cystine, and hydroxy-proline were minor constituents (0.0–2.6%). Glucose was the predominant sugar in all cultures, ranging from 23.0% (50 μE·m^?2·s^?1) to 45.0% (100 μE·m^?2·s^?1) of total polysaccharide. No correlation was found between the proportion of any of the sugars and irradiance. The proportions of the lipid class components and fatty acids showed little change with irradiance. The main fatty acids were 14:0, 16:0, 16:1(n-7), 18:1(n-9), 18:3(n-3). 18:4(n-3), 18:5(n-3), and 22:6(n-3). Proportions of 22: 6(n-3) increased, whereas l8:3(n-3). 18:3(n-6). and 18:4(n-3) decreased, with increasing irradiance. Pigment concentrations were highest in cultures grown at 50 μE·m^?2·s^?1, except for fucoxanthin and diadinoxanthin (100 μE·m^?2·s^?1). The concentrations of accessory pigments correlated with chlorophyll a, which decreased in concentration with increasing irradiance. On the basts of biochemical composition, an irradiance of 100 μE·m^?1·s^?1 (12:12 h L:D cycle)for the culture of Isochrysis sp. (clone T-ISO) may provide optimal nutritional value for maricultured animals, although feeding trials are now necessary to substantiate this.     

PHOTOSYNTHESIS AND CELL DIVISION BY ANTARCTIC MICROALGAE: COMPARISON OF BENTHIC,PLANKTONIC AND ICE ALGAE1

Irradiance-dependent rates of photosynthesis and cell division of six species of microalgae isolated from the benthos, plankton and sea ice microbial community in McMurdo Sound, Antarctica were compared. Microalgae isolated from different photic environments had distinct photosynthetic and growth characteristics. For benthic and ice algae, photosynthesis saturated at 6 to 20 μE.m^?2.s^?1 and was photoinhibited at 10 to 80 μE.m^?2.s^?1 while for the planktonic algae, saturation irradiances were up to 13 times higher and photoinhibition was not detected. The slope of the light-limited portion of the P-I relationship was up to 50 times greater for the benthic algae than for either the ice or planktonic algae suggesting that benthic algae used the low irradiances more efficiently for carbon uptake. Cell division was dependent on the incubation irradiance for all but one microalga examined. The dependence of division rates on irradiance was however much smaller than for carbon uptake, suggesting that cell division buffers the influence of short term variations of irradiance on cellular metabolism.     

Effects of light intensity and nutrients on gametophytes and gametogenesis of Lessonia nigrescens Bory (Phaeophyta)

Meiospores of Lessonia nigrescens Bory collected in Central Chile, were cultivated under different light and nutrient conditions to test for their influence on the development of gametophytes and gametogenesis. The microscopic stages of the life cycle are described. Under 45 μE·m^?2 ·s^?1 (12 h daily) and sea water enriched with nitrate and phosphate, development of few-celled gametophytes, gametogenesis and sporophyte differentiation occur within 35 days. At 10 μE ·m^?2 · s^?1, most gametophytes are multicellular and remain vegetative up to 65 days. Only enriched sea water or enriched sea water with double concentrations of phosphate lead to fertile few-celled gametophytes under 45 μE · m^?2 · s^?1. The addition of double concentrations of nitrate allows for the development of few-celled gametophytes but inhibits gametogenesis. The average gametophyte size progressively decreases when increasing amounts of nitrate and phosphate are added to the media.     

EFFECT OF TEMPERATURE,LIGHT AND pH ON GROWTH,PHOTOSYNTHESIS AND RESPIRATION OF THE DINOFLAGELLATE PERIDINIUM CINCTUM FA. WESTII IN LABORATORY CULTURES1

Optimum light, temperature, and pH conditions for growth, photosynthetic, and respiratory activities of Peridinium cinctum fa. westii (Lemm.) Lef were investigated by using axenic clones in batch cultures. The results are discussed and compared with data from Lake Kinneret (Israel) where it produces heavy blooms in spring. Highest biomass development and growth rates occurred at ca. 23° C and ≥50 μE· m^?2·s¹ of fluorescent light with energy peaks at 440–575 and 665 nm. Photosynthetic oxygen release was more efficient in filtered light of blue (BG 12) and red (RG 2) than in green (VG 9) qualities. Photosynthetic oxygen production occurred at temperatures ranging from 5° to 32° C in white fluorescent light from 10 to 105 μE·m^?2·s^?1 with a gross maximum value of 1500 × 10^?12 g·cell^?1·h^?1 at the highest irradiance. The average respiration amounted to ca. 12% of the gross production and reached a maximum value of ca. 270·10^?12 g·cell^?1·h^?1 at 31° C. A comparison of photosynthetic and respiratory Q₁₀-values showed that in the upper temperature range the increase in gross production was only a third of the corresponding increase in respiration, although the gross production was at maximum. Short intermittent periods of dark (>7 min) before high light exposures from a halogen lamp greatly increased oxygen production. Depending on the physiological status of the alga, light saturation values were reached at 500–1000 μE·m^?2·s^?1 of halogen light with compensation points at 20–40 μE·m^?2·s^?1 and I_k-values at 100–200 μE·m^?2·s^?1. The corresponding values in fluorescent light in which it was cultured and adapted, were 25 to 75% lower indicating the ability of the alga to efficiently utilize varying light conditions, if the adaptation time is sufficient. Carbon fixation was most efficient at ca. pH 7, but the growth rates and biomass development were highest at pH 8.3.     

PHOTOSYNTHESIS IN AN EXTREME SHADE ENVIRONMENT: BENTHIC MICROBIAL MATS FROM LAKE HOARE, A PERMANENTLY ICE-COVERED ANTARCTIC LAKE

We investigated the composition of benthic microbial mats in permanently ice-covered Lake Hoare, Antarctica, and their irradiance vs. photosynthetic oxygen exchange relationships. Mats could be subdivided into three distinct depth zones: a seasonally ice-free “moat” zone and two under-ice zones. The upper under-ice zone extended from below the 3.5 m thick ice to approximately 13 m and the lower from below 13 m to 22 m. Moat mats were acclimated to the high irradiance they experienced during summer. They contained photoprotective pigments, predominantly those characteristic of cyanobacteria, and had high compensation and saturating irradiances (E_c and E_k) of 75 and 130 μmol photons·m⁻²·s⁻¹, respectively. The moat mats used light inefficiently. The upper under-ice community contained both cyanobacteria and diatoms. Within this zone, biomass (as pigments) increased with increasing depth, reaching a maximum at 10 m. Phycoerythrin was abundant in this zone, with shade acclimation and efficiency of utilization of incident light increasing with depth to a maximum of 0.06 mol C fixed·mol⁻¹ incident photons under light-limiting conditions. Precipitation of inorganic carbon as calcite was associated with this community, representing up to 50% of the carbon sequestered into the sediment. The lower under-ice zone was characterized by a decline in pigment concentrations with depth and an increasing prevalence of diatoms. Photosynthesis in this community was highly shade acclimated and efficient, with E_c and E_k below 0.5 μmol·m⁻²·s⁻¹ and 2 μmol·m⁻²·s⁻¹, respectively, and maximum yields of 0.04 mol C fixed·mol⁻¹ incident quanta. Carbon uptake in situ by both under-ice and moat mats was estimated at up to 100 and 140 mg·m⁻²·day⁻¹, based on the photosynthesis–irradiance curves, incident irradiance, and light attenuation by ice and the water column.     

Relationships between irradiance and photosynthesis for marine benthic green algae (Chlorophyta) of differing morphologies

Photosynthesis-irradiance relationships were determined in the field for five species of littoral and shallow sublittoral marine benthic green algae (Chlorophyta) of differing morphologies. Each species exhibited a linear increase in photosynthetic rate with increasing irradiance up to a maximum light-saturated value. Full sunlight (1405 to 1956 μE·m^?2·s^?1) inhibited photosynthesis of all species except the thick, optically dense, Codium fragile (Sur.) Har. Compensation irradiances ranged from 6.1 μE·m^?2·s^?1 for Enteromorpha intestinalis (L.) Link to 11.4 μE·m^?2·s^?1 for Ulva lobata (Kütz) S. & G. and did not reveal a consistent relationship to seaweed morphology. Saturation irradiances were determined statistically (I_k) and visually from graphical plots. with the latter technique resulting in values three to eight times higher and different comparative rankings of species than the former. I_k saturation irradiances were highest for Chaetomorpha linum (Müll.) Kütz. (81.9 μE·m^?2·s^?1) and lowest for Codium fragile (49.6 μE·m^?2·s^?1) and did not reveal a relationship with seaweed morphology. Regression equations describing light-limited photosynthetic rates and the relative magnitudes of the maximal net photosynthetic responses both strongly suggested a relationship with seaweed morphology. Highest net photosynthetic rates were obtained for the thin, sheet-like algae Ulva lobata (9.2 mg C·g dry wt^?1·h^?1), U. rigida C. Ag. (6.5 mg C·g dry wt^?1·h^?1) and the tubular form, Enteromorpha intestinalis (7.3 mg C·g dry wt^?1·h^?1), while lowest rates occurred for Codium fragile (0.9 mg C·g dry wt^?1·h^?1). Similarly, steepest light-limited slopes were found for the algae of simpler morphology, while the most gradual slope was determined for Codium fragile, the alga with greatest thallus complexity.     

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.     

IMPORTANCE OF PHYSICAL VARIABLES ON THE SEASONAL DYNAMICS OF EPILITHIC ALGAE IN A HIGHLY SHADED CANYON STREAM1

The seasonal abundance of epilithic algae was correlated with major physico-chemical parameters in a first-order, heavily shaded stream in northern Arizona. Diatoms made up over 85%, by numerical abundance, of the epilithon community Light energy, water temperature, and stream discharge were most highly correlated with seasonal abundance of epilithic diatom taxa when analyzed with stepwise multiple regression. None of the chemical variables measured in the study (NO₃-N, O-PO₄, SiO₂, including PH) was found to be significantly correlated with the seasonal community structure of epilithic diatoms. Total diatom cell densities showed a significant negative correlation to stream bed light energy. Likewise, total diatom cell densities along a transect in the stream bed showed a negative correlation to current velocity during those months when base flow was low and stable, and current velocity was ≤25 cm·sec-¹. Most diatom taxa had highest cell densities at temperatures < 16°C and at daily mean stream bed light levels < 400 μE·m^?2·s^?1. Highest cell densities of green algae occurred at temperatures between 6–16°C and at daily mean stream bed light levels of > 400 μE·m^?2·s^?1. Blue-green algae (cyanobacteria) grew best at the highest recorded water temperatures and daily mean stream bed light energy (16–20°C and 900–1200 μE·m^?2·s^?1). Abrupt increases in NO₃-N coincided with a brief pulse of Nostoc pruniforme colonies during June, and leaf drop from Alnus oblongifolia during October.     

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.     

EFFECTS OF SUBSTRATE RELIEF ON THE DISTRIBUTION OF PERIPHYTON IN LABORATORY STREAMS. II. INTERACTIONS WITH IRRADIANCE1

Laboratory streams were used in a 42-day experiment designed to investigate how the spatial and temporal distribution of lotic periphyton created by current flow over cobble-size substrates is a affected by irradiance. The streams contained 22.5 × 22.5 × 4 cm substrate blocks and were exposed to either 385, 90 or 20 μE·m^?2·s^?1. We monitored periphyton succession in fast current regimes on top of blocks and in slower current regimes on surfaces recessed between blocks. The absolute differences in AFDW algal biomass between top and recessed substrates were significantly affected by irradiance and time. At the end of the experiment, biomass in streams exposed to 385 μE·m^?2·s^?1. was approximately 2 and 8 times greater than in streams exposed to 90 and 20 μE·m^?2·s^?1, respectively. Differences in biomass were greater between irradiance levels than between top and recessed substrates within an irradiance level. Irradiance also had a greater effect than current regime on the taxonomic composition of assemblages. Oscillatoria agardhii Gomont and Navicula minima Grun. characterized assemblages at 20 μE·m^?2·s^?1, whereas Fragilaria vaucheriae (Kütz.), Nitzschia oregona Sov., Navicula arvensis Hust. and Stigeoclonium tenue (Ag.) Kütz. were more abundant at the two higher irradiances. Detrended correspondence analysis indicated that the rate of succession was relatively high for assemblages at high irradiance and in the slow current regimes between blocks. The results suggested that in natural streams, periphyton patches produced by large differences in irradiance should have a greater effect on periphyton heterogeneity than substrate-induced patches. Moreover, the heterogeneity of algal patches produced by hydrologic differences over a substrate is constrained by irradiance level.     

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.     

LIGHT‐INDUCED MOTILE RESPONSES OF THE ESTUARINE BENTHIC DIATOMS NAVICULA PERMINUTA AND CYLINDROTHECA CLOSTERIUM (BACILLARIOPHYCEAE)1

Motility of estuarine epipelic (mud‐inhabiting) diatoms is an important adaptation to living in biofilms present within fine sediments. Motility allows cells to migrate within the photic zone in response to a wide range of environmental stimuli. The motile responses of two species of benthic diatoms to photon fluence rates and spectral quality were investigated. Cultures of Navicula perminuta (Grunow) in van Heurck and Cylindrotheca closterium (Ehrenb.) J. C. Lewin et Reimann both exhibited photoaccumulation at ～200 μmol · m^?2 · s^?1 and photodispersal from photon flux densities (PFDs) of ～15 μmol · m^?2 · s^?1. Photokinesis (changing cell speed) contributed toward photodispersal for both species, and red light (λ = 681–691 nm) was most effective at inducing this process. N. perminuta showed a phototactic (directional) response, with active movement in response to a light gradient. Although this response was exhibited in white light, these directional responses were only elicited by wavelengths from 430 to 510 nm. In contrast, C. closterium did not exhibit phototaxis under any light conditions used in this study. Motile benthic diatoms thus exhibit complex and sophisticated responses to light quantity and quality, involving combinations of photokinesis and phototaxis, which can contribute toward explaining the patterns of large‐scale cell movements observed in natural estuarine biofilms.     

EFFECTS OF IRRADIANCE AND GRAZING ON LOTIC ALGAL ASSEMBLAGES1

A laboratory experiment was conducted for 75 days to examine how irradiance levels and grazing influence algal biomass and community structure. Twelve laboratory streams were used for experimental analyses, with four channels exposed to one of three irradiance levels (15, 100, or 400 μE·m^?2·s^?1). Three of the four stream at each light level were stocked with the snail Juga silicula (250·m^?2), leaving one stream at each light level without snails. Grazed stream exposed to low light levels developed low amounts of algal biomass (<2 g AFDW·m^?2) and were dominated by adnately attached diatoms. Mean algal biomass increased over time in the grazed streams exposed to intermediate light; by day 75, these streams were characterized by moderate algal biomasses (30-40 g AFDW·m^?2) and filamentous chlorophytes. Algal assemblages in high light, grazed channels had high levels of biomass at day 43 (70 g AFDW·m^?2) that declined to 30 g AFDW·m^?2at day 75 and were dominated by chlorophytes. Among ungrazed streams, algal biomass at day 75 was relatively low in the low light streams (<7g AFDW·m^?2) and was dominated by adnately attached diatoms. Ungrazed streams exposed to intermediate and high light levels had moderate biomasses (23 and 19 g AFDW·m^?2, respectively) and were dominated by chlorophytes and large diatoms. Grazing appeared both to delay and alter successional trajectories of algal assemblages, with alterations most noticeable during early seral stages at intermediate and high light levels. Grazing had the least effect on successional trajectories at low light.     

LIGHT,TEMPERATURE AND PHOTOPERIOD AS FACTORS CONTROLLING REPRODUCTION IN ULOTHRIX ZONATA (ULVOPHYCEAE)1

In many lakes in the northern United States and Canada the filamentous green alga Ulothrix zonata (Weber and Mohr) Kütz grows abundantly in early spring in shallow waters. Asexual reproduction occurs by formation of quadriflagellate zoospores which disrupt, the integrity of the cells upon release causing the filament to disintegrate. Study of the effects of 100 different combinations of irradiance, temperature and photoperiod revealed that zoospore formation is favored by high temperatures near 20°C, high light levels of 520 μE·m^?2·s^?1 and photoperiods of either short day (8:16 h light-dark) or long day cycles (16:8 h light-dark). Zoospore formation is minimal under conditions of low temperature (5°C), low irradiance (32.5 μE·m^?2·s^?1) and neutral day-lengths (12:12 h light-dark). These observations explain the decline in U. zonata biomass when water temperatures rise above 10° C. The combined effect of rising water temperatures and increasing daylengths causes progressively more filaments to switch from vegetable growth to zoospore production resulting in an increasing loss of biomass.