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.     

Intraspecific egg size variation and sperm limitation in the broadcast spawning bivalve Macoma balthica

Bottom-ice algae within Antarctic sea ice were examined using chlorophyll fluorescence imaging. The detailed structure of the bottom-ice algal community growing in the platelet and congelation layers of solid pieces of sea ice was evident for the first time in chlorophyll imaging mode. Strands of fluorescence representing algal cells were clearly visible growing upward into brine channels in a fine network. Images of effective quantum yield (Ф_PSII) revealed that the Ф_PSII of algae embedded in the sea ice was approximately 0.5. Furthermore, Ф_PSII decreased slightly with distance from the ice-water interface.The response of Antarctic sea ice algae to changes in irradiance and salinity, and the effects of slowly warming and melting the ice block sample were examined using this system. The Ф_PSII of bottom-ice algae decreased as irradiance increased and salinities decreased. Bottom-ice algae appear to be most vulnerable to changes in their environment during the melting process of the ice, and this suggests that algae from this region of the ice may not be able to cope with the stress of melting during summer.Chlorophyll fluorescence imaging provides unprecedented imagery of chlorophyll distribution in sea ice and allows measurement of the responses of sea ice algae to environmental stresses with minimal disruption to their physical habitat. The results obtained with this method are comparable to those obtained with algae that have been melted into liquid culture and this indicates that previous melting protocols reveal meaningful data. In this chlorophyll imaging study, rapid light curves did not saturate and this may prevent further use of this configuration.     

Small scale vertical gradients of Arctic ice algal photophysiological properties

Photosynthetic parameters of phytoplankton and sea ice algae from landfast sea ice of the Chukchi Sea off Point Barrow, Alaska, were assessed in spring 2005 and winter through spring 2006 using Pulse Amplitude Modulated (PAM) fluorometry including estimates of maximum quantum efficiency (F _v/F _m), maximum relative electron transport rate (rETR_max), photosynthetic efficiency (α), and the photoadaptive index (E _k). The use of centrifuged brine samples allowed to document vertical gradients in ice algal acclimation with 5 cm vertical resolution for the first time. Bottom ice algae (0–5 cm from ice–water interface) expressed low F _v/F _m (0.331–0.426) and low α (0.098–0.130 (μmol photons m⁻²s⁻¹)⁻¹) in December. F _v/F _m and α increased in March and May (0.468–0.588 and 0.141–0.438 (μmol photons m⁻²s⁻¹)⁻¹, respectively) indicating increased photosynthetic activity. In addition, increases in rETR_max (3.3–16.4 a.u.) and E _k (20–88 μmol photons m⁻² s⁻¹) from December to May illustrates a higher potential for primary productivity as communities become better acclimated to under-ice light conditions. In conclusion, photosynthetic performance by ice algae (as assessed by PAM fluorometry) was tightly linked to sea ice salinity, temperature, and inorganic nutrient concentrations (mainly nitrogen).     

The effects of desiccation and salinity gradients on the PSII photochemical efficiency of an intertidal brown alga,Sargassum fusiforme from Kagoshima,Japan

The responses of photochemical efficiency to desiccation and salinity gradients in an intertidal edible brown macroalga, Sargassum fusiforme (Harvey) Setchell (Sargassaceae, Fucales), were determined using a pulse amplitude modulation (PAM)-chlorophyll fluorometer. The effective quantum yields (ΔF/F_m'; = Φ_PSII) of photosystem II (PSII) dropped to zero after 360-min aerial exposure under low irradiance (20 μmol photons m⁻² s⁻¹) and 120-min exposure under high irradiance (700 μmol photons m⁻² s⁻¹) for this species at 20°C and 50% relative humidity. Under these conditions, ΔF/F_m' failed to recover to initial levels even after 1-day rehydration in seawater. In general, ΔF/F_m' decreased as desiccation reduced the absolute water content (AWC, %). Nevertheless, when AWC was above ca. 20%, ΔF/F_m' was mostly restored to initial levels after 1-day rehydration in seawater, suggesting strong tolerance to dehydration. Furthermore, S. fusiforme appeared to tolerate a broad range of salinity (i.e. 15–50 psu) during six days of culture; however, ΔF/F_m' declined when salinity was <10 and 60 psu. Strong tolerance to dehydration and salinity stress likely provides S. fusiforme an advantage that allows it to flourish in the intertidal habitat.     

FLUORESCENCE-BASED MAXIMAL QUANTUM YIELD FOR PSII AS A DIAGNOSTIC OF NUTRIENT STRESS

In biological oceanography, it has been widely accepted that the maximum quantum yield of photosynthesis is influenced by nutrient stress. A closely related parameter, the maximum quantum yield for stable charge separation of PSII, (φ _PSII )_m, can be estimated by measuring the increase in fluorescence yield from dark-adapted minimal fluorescence (F_o) to maximal fluorescence (F_m) associated with the closing of photosynthetic reaction centers with saturating light or with a photosynthetic inhibitor such as 3′-(3,4-dichlorophenyl)-1′,1′-dimethyl urea (DCMU). The ratio F_v/F_m (= (F_m− F_o)/F_m) is thus used as a diagnostic of nutrient stress. Published results indicate that F_v/F_m is depressed for nutrient-stressed phytoplankton, both during nutrient starvation (unbalanced growth) and acclimated nutrient limitation (steady-state or balanced growth). In contrast to published results, fluorescence measurements from our laboratory indicate that F_v/F_m is high and insensitive to nutrient limitation for cultures in steady state under a wide range of relative growth rates and irradiance levels. This discrepancy between results could be attributed to differences in measurement systems or to differences in growth conditions. To resolve the uncertainty about F_v/F_m as a diagnostic of nutrient stress, we grew the neritic diatom Thalassiosira pseudonana (Hustedt) Hasle et Heimdal under nutrient-replete and nutrient-stressed conditions, using replicate semicontinuous, batch, and continuous cultures. F_v/F_m was determined using a conventional fluorometer and DCMU and with a pulse amplitude modulated (PAM) fluorometer. Reduction of excitation irradiance in the conventional fluorometer eliminated overestimation of F_o in the DCMU methodology for cultures grown at lower light levels, and for a large range of growth conditions there was a strong correlation between the measurements of F_v/F_m with DCMU and PAM (r² = 0.77, n = 460). Consistent with the literature, nutrient-replete cultures showed consistently high F_v/F_m (∼0.65), independent of growth irradiance. Under nutrient-starved (batch culture and perturbed steady state) conditions, F_v/F_m was significantly correlated to time without the limiting nutrient and to nutrient-limited growth rate before starvation. In contrast to published results, our continuous culture experiments showed that F_v/F_m was not a good measure of nutrient limitation under balanced growth conditions and remained constant (∼0.65) and independent of nutrient-limited growth rate under different irradiance levels. Because variable fluorescence can only be used as a diagnostic for nutrient-starved unbalanced growth conditions, a robust measure of nutrient stressed oceanic waters is still required.     

MICROALGAL LIGHT-HARVESTING IN EXTREME LOW-LIGHT ENVIRONMENTS IN MCMURDO SOUND, ANTARCTICA1

Microalgal pigment composition, photosynthetic characteristics, single-cell absorption efficiency (Qa_(λ)) spectra, and fluorescence-excitation (FE) spectra were determined for platelet ice and benthic communities underlying fast ice in Mc Murdo Sound, Antarctica, during austral spring 1988. Measurements of spectral irradiance (E_(λ)) and photosynthetically active radiation (PAR) as well as samples for particulate absorption measurements were taken directly under the congelation ice, within the platelet layer, as profiles vertically through the water column, and at the benihic surface. Light attenuation by.sea ice, algal pigments, and particulates reduced PAR reaching the platelet ice layer to 3%(9–33 fimol photons m-²-^?s-¹) of surface values and narrowed its spectral distribution to a band between 400 and 580 nm. Attenuation by the water column further reduced PAR reaching the sea floor (28–m depth) to 0.05% of surface levels (< 1 μmol photons m-² s-¹), with a spectral distribution dominated by 470–580–nm wavelengths. The photoadaptive index (I) for platelet ice algae (5.9–12.6 μmol photons m-^2.s-¹) was similar to ambient PAR, indicating that algae had acclimated to their light environment (i.e. the algae were light-replete). Maximum Qa_(λ) at the blue absorption peak (440 nm) was 0.63, and enhanced absorption was observed from 460–500 nm and was consistent with observed high cellular chlorophyll (chi) c:chl a and fucoxanthin: chl a molar ratios (0.4 and 1.2, respectively). Benthic algae were light-limited despite the maintenance of very low I_k values (4–11 μmol photons.m-^2.s-¹). Extremely high fucoxanthin: chi a ratios (1.6) in benthic algae produced enhanced green light absorption, resulting in a high degree of complementation between algal absorption and ambient spectral irradiance. Qa_(λ) values for benthic algae were maximal (0.9) between 400 and 510 nm but remained >0.35 even at absorption minima. Strong spectral flattening, a characteristic of intense pigment packaging, was also apparent in the Qa_(λ) spectra for benthic algae. FE and Qa_(λ) spectra were similar in shape for platelet ice algae, indicating that the efficiency at which absorbed energy was transferred to photosystem II (PSII) was independent of wavelength. Fluorescence emission by benthic algae was greatest for the 500–560–nm excitation wavelengths, suggesting that most energy absorbed by accessory pigments was transferred to PSII. These results suggest that under ice algae employ complementary pigmentation and maximize absorption efficiency as adaptive strategies to low-light stress. Regulating the distribution of absorbed energy between PSI and PSII may be an adaptive response to the restricted spectral distribution of irradiance.     

Photosynthetic activity in two heteromorphic life‐history stages of a freshwater red alga,Thorea gaudichaudii (Thoreales) from Japan,in response to an irradiance and temperature gradient

We determined the effect of irradiance and temperature on the photosynthesis of two heteromorphic life‐history stages of an endangered freshwater red alga, Thorea gaudichaudii (Thoreales) by laboratory and field measurements. Net oxygenic photosynthesis–irradiance models of macroscopic and microscopic life‐history stages revealed similar low irradiance‐adapted responses, with a compensation irradiance (E_c) of 6.71 and 2.56 μmol photons m^?2 s^?1 (4.30–9.13 and 0.13–7.19, 95% Bayesian prediction interval, BPI) and saturating irradiance (E_k) of 26.6 and 30.0 μmol photons m^?2 s^?1 (19.0–37.4 and 12.1–63.0, BPI), respectively. A temperature‐dependent model of net photosynthesis and dark respiration in macroscopic and microscopic stages also showed similar temperature responses, and the gross photosynthetic rate (GP_max), 3.54 and 6.34 μg O₂ g_ww^?1 min^?1 (3.10–3.99 and 5.31–8.21, BPI), was highest at 32.1 and 35.7°C (29.8–34.0 and 29.5–48.6, BPI). The maximum quantum yields (F _v/F _m) in macroscopic and microscopic stages were also similar in response with respect to temperature; however, it was somewhat steady at low temperatures with the highest value of 0.54 and 0.62 (0.54–0.55 and 0.61–0.63, BPI) at 17.8 and 15.0°C (16.7–18.8 and 12.3–17.1, BPI). The effective quantum yield (Φ _PSII) in macroscopic and microscopic stages was also negatively correlated with irradiance, which decreased after 12 h of continuous exposure to 50 (low) and 1000 (high) μmol photons m^?2 s^?1 at 12 and 22°C. Large declines of Φ _PSII and subsequent failure of F _v/F _m recovery were particularly enhanced at high irradiance, signifying photoinhibition. Diurnal change of Φ _PSII and incident irradiance of the macroscopic stage under the field measurement revealed the midday depression of Φ _PSII; however, there was little direct sunlight due to shading by the trees, and algae were occurring in the shaded locations in the freshwater spring.     

Physiological ecology of photosynthesis in Prasiola stipitata (Trebouxiophyceae) from the Bay of Fundy,Canada

The physiological ecology of Prasiola stipitata was examined in situ from two supralittoral sites in the Bay of Fundy (Nova Scotian, Canada) during November 2011, when the population was undergoing major expansion. Photosynthetic parameters (effective quantum yield, Φ_PSII, maximum quantum yield, F_v/F_m, and relative electron transport rate, rETR) were evaluated using chlorophyll fluorescence of PSII. A largely shaded and continuously moist population showed no change in Φ_PSII from one hour after sunrise to sunset in which natural irradiance varied between 3 and 300 μmol photons m^?2 s^?1. High irradiance (up to 1800 μmol photons m^?2 s^?1) had no apparent negative impacts on either quantum yield or rETR, but high desiccation in the field reduced quantum yield to almost zero. When thalli were brought into the laboratory, no change in F_v/F_m was observed up to 60% dehydration; however, there was a steep decline in F_v/F_m between 60% and 85% dehydration. Thalli showed complete recovery of F_v/F_m within one hour of reimmersion in seawater after 2 days of desiccation. After 15 days of desiccation full recovery required 24 h and after 30 days of desiccation thalli showed only partial recovery. These observations confirm the adaptation to photosynthesis in high irradiances and the rapid recovery following extreme desiccation observed in other Prasiola species.     

Quantification of algal biofilms colonising building materials: chlorophyll a measured by PAM-fluorometry as a biomass parameter

Abstract

The aim of this study was to quantify algal colonisation on anthropogenic surfaces (viz. building facades and roof tiles) using chlorophyll a (chl a) as a specific biomarker. Chl a was estimated as the initial fluorescence F₀ of ‘dark adapted’ algae using a pulse-modulated fluorometer (PAM-2000). Four isolates of aeroterrestrial green algae and one aquatic isolate were included in this study. The chl a concentration and F₀ showed an exponential relationship in the tested range between 0 and 400 mg chl a m^?2. The relationship was linear at chl a concentrations <20 mg m^?2. Exponential and linear models are presented for the single isolates with large coefficients of determination (exponential: r² > 0.94, linear: r² > 0.92). The specific power of this fluorometric method is the detection of initial algal colonisation on surfaces in thin or young biofilms down to 3.5 mg chl a m^?2, which corresponds to an abundances of the investigated isolates between 0.2 and 1.5 million cells cm^?2.     

The effect of temperature,light-spectrum,desiccation and salinity gradients on the photosynthetic performance of a subtidal brown alga,Sargassum macrocarpum,from Japan

The effect of temperature, light-spectrum, desiccation and salinity gradients on the photosynthesis of a Japanese subtidal brown alga, Sargassum macrocarpum (Fucales), was determined using a pulse amplitude modulation-chlorophyll fluorometer and dissolved oxygen sensors. Temperature responses of the maximum (F_v/F_m in darkness) and effective (ΔF/F_m′ at 50 μmol photons m⁻² s⁻¹; = Φ_PSII) quantum yields during 6-day culture (4–36°C) remained high at 12–28°C, but decreased at higher temperatures. Nevertheless, ΔF/F_m′ also dropped at temperatures below 8°C, suggesting light sensitivity under chilling temperatures because F_v/F_m remained high. Photosynthesis–irradiance responses at 24°C under red (660 nm), green (525 nm), blue (450 nm) and white light (metal halide lamp) showed that maximum net photosynthesis under blue and white light was greater than under red and green light, indicating the sensitivity and photosynthetic availability of blue light in the subtidal light environment. In the desiccation experiment, samples under aerial exposure of up to 8 h under dim-light at 24°C and 50% humidity showed that ΔF/F_m′ quickly declined after more than 45 min of emersion; furthermore, ΔF/F_m′ also failed to recover to initial levels even after 1 day of rehydration in seawater. Under the emersion state, the ΔF/F_m′ remained high when the relative water content (RWC) was greater than 50%; in contrast, it quickly dropped when the RWC was less than 50%. When the RWC was reduced below 50%, ΔF/F_m′ did not return to initial levels, regardless of subsequent re-hydration, suggesting a low capacity of photosynthesis to recover from desiccation. The stenohaline response of photosynthesis under 3-day culture is evident, given that ΔF/F_m′ declined when salinity was beyond 20–40 psu. Adaptation to subtidal environments in temperate waters of Japan can be linked to these traits.     

FLUORESCENCE INDUCTION AND PHOTOSYNTHETIC RESPONSES OF ARCTIC ICE ALGAE TO SAMPLE TREATMENT AND SALINITY1

The measurement of Photosynthetic rates of algae growing on the undersurface of 1. 7 m thick ice in the Canadian Arctic (Resolute Passage. N.W.T.) presents several problems. During the preparation of samples for physiological measurements, the ice algae may he exposed to salinity and temperature shocks. Fluorescence induction (the rise in in vivo Chl a fluorescence intensity during a period of millineconds) and photosynthesis-irradiance (PI) experiments examined the potential effects of salinity and temperature on the physiology of ice algae. Experimental suspensions were routinely prepared by scraping one part ire crystals (11–14%₀ salinity) and attached algae from the bottom ice into four parts filtered seawater (32%₀ salinity). giving a final salinity of 28–31%₀. Post-dilution of melted ice scrapings with seawater suppressed photosynthetic ¹⁴C-fixation and decreased A_DCMU (the area above the fluorescence induction curve measured in the presence of the inhibitor DCMC: an estimate of photosynthetic capacity) by a factor of 3–16. due to the low salinity of the melted ice scrapings. Fluorescence induction and PI experiments showed that the ice algae had a salinity optimum near 30%₀, close to the ambient seawater salinity, Experiments in which the Chl a concentration was manipulated showed that A_DCMU, P^a_m (Chl a-normalized rate of photosynthesis at light saturation), and a (photosynthetic efficiency) declined with increasing Chl a concentration. Ice algae tolerated heating (l.5°C-min^-1) up to 17° C, above which A_DCMU’decreased with sample temperature.     

Linking ice structure and microscale variability of algal biomass in Arctic first-year sea ice using an in situ photographic technique

Microscale photographs were taken of the ice bottom to examine linkages of algal chlorophyll a (chl a) biomass distribution with bottom ice features in thick Arctic first-year sea ice during a spring field program which took place from May 5 to 21, 2003. The photographic technique developed in this paper has resulted in the first in situ observations of microscale variability in bottom ice algae distribution in Arctic first-year sea ice in relation to ice morphology. Observations of brine channel diameter (1.65–2.68 mm) and number density (5.33–10.35 per 100 cm²) showed that the number of these channels at the bottom of thick first-year sea ice may be greater than previously measured on extracted ice samples. A variogram analysis showed that over areas of low chl a biomass (≤20.7 mg chl a m⁻²), patchiness in bottom ice chl a biomass was at the scale of brine layer spacing and small brine channels (∼1–3 mm). Over areas of high chl a biomass (≥34.6 mg chl a m⁻²), patchiness in biomass was related to the spacing of larger brine channels on the ice bottom (∼10–26 mm). Brine layers and channels are thought to provide microscale maxima of light, nutrient replenishment and space availability which would explain the small scale patchiness over areas of low algal biomass. However, ice melt and erosion near brine channels may play a more important role in areas with high algal biomass and low snow cover.     

Succession and physiological health of freshwater microalgal fouling in a Tasmanian hydropower canal

Freshwater microalgal biofouling in hydropower canals in Tarraleah, Tasmania, is dominated by a single diatom species, Gomphonema tarraleahae. The microfouling community is under investigation with the aim of reducing its impact on electricity generation. Species succession was investigated using removable glass slides. Fouled slides were examined microscopically and for chlorophyll a biomass. Chl a biomass increased steeply after 8 weeks (0.09–0.87 mg m^?2), but increased much earlier on slides surrounded by a biofouled inoculum. Succession began with low profile diatoms such as Tabellaria flocculosa, progressing to stalked diatoms such as Gomphonema spp. and Cymbella aspera. Few chlorophytes and no filamentous algae were present. Pulse amplitude modulated fluorometry was used to measure the physiological health of fouling on the canal wall. Maximum quantum yield (F _v/F _m) measurements were consistently <0.18, indicating that the fouling mat consisted of dead or dying algae. The succession and physiological health of cells in the fouling community has broad implications for mitigation techniques used.     

USE OF PULSE‐AMPLITUDE‐MODULATED FLUORESCENCE TO ASSESS THE PHYSIOLOGICAL STATUS OF CLADOPHORA SP. ALONG A WATER QUALITY GRADIENT1

This study investigated the application of pulse‐amplitude‐modulated (PAM) fluorometry as a rapid assessment of benthic macroalgal physiological status. Maximum quantum efficiency (F_v/F_m), dark–light induction curves, and rapid fluorescence light‐response curves (RLC) were measured on the filamentous macroalgal Cladophora sp. from Lake Ontario on 5 d at 16 sites spanning a gradient of light and nutrient supply. For Cladophora sp. growing in situ, light limitation was assessed by comparing average daily irradiance with the light utilization efficiency parameter (α) derived from RLCs. In this study, there was a nonlinear relationship between F_v/F_m and the degree of P limitation in macroalgae. However, only light‐saturated Cladophora sp. showed a significant positive linear relationship between F_v/F_m and P nutrient status. The absence of this relationship among light‐limited algae indicates that their photosynthetic rate would be stimulated by increased water clarity, and not by increased P supply. PAM fluorescence measures were successfully able to identify light‐saturated macroalgae and, among these, assess the degree to which they were nutrient limited. These results enable us to test hypotheses arising from numeric models predicting the impact of changes in light penetration and nutrient supply on benthic primary production.     

Photosynthetic activity including the thermal‐ and chilling‐light sensitivities of a temperate Japanese brown alga Sargassum macrocarpum

The effects of irradiance, temperature, thermal‐ and chilling‐light sensitivities on the photosynthesis of a temperate alga, Sargassum macrocarpum (Fucales) were determined by a pulse amplitude modulation (PAM)‐chlorophyll fluorometer and dissolved oxygen sensors. Oxygenic photosynthesis–irradiance curves at 8, 20, and 28°C revealed that the maximum net photosynthetic rates (NP _max) and saturation irradiance were highest at 28°C, and lowest at 8°C. Gross photosynthesis and dark respiration determined over a range of temperatures (8–36°C) at 300 μmol photons m^?2 s^?1 revealed that the maximum gross photosynthetic rate (GP_max) occurred at 27.8°C, which is consistent with the highest seawater temperature in the southern distributional limit of this species in Japan. Additionally, the maximum quantum yields of photosystem II (F _v/F _m) during the 72‐h temperature exposures were stable at 8–28°C, but suddenly dropped to zero at higher temperatures, indicative of PSII deactivation. Continuous exposure (12 h) to irradiance of 200 (low) and 1000 (high) μmol photons m^?2 s^?1 at 8, 20, and 28°C revealed greater declines in their effective quantum yields (Φ _PSII) under high irradiance. While Φ _PSII under low irradiance were very similar with the initial F _v/F _m under 20 and 28°C, values rapidly decreased with exposure duration at 8°C. At this temperature, F _v/F _m did not recover to initial values even after 12 h of dark acclimation. Final F _v/F _m of alga at 28°C under high irradiance treatment also did not recover, suggesting its sensitivity to photoinhibition at both low and high temperatures. These photosynthetic characteristics reflect both the adaptation of the species to the general environmental conditions, and its ability to acclimate to seasonal changes in seawater temperature within their geographical range of distribution.     

Subsurface chlorophyll maximum (SCM) location and extension in the water column as governed by a density interface in the strongly stratified Kattegat estuary

The aims of the study were to analyse the relations between the physics of a water column and the location of the subsurface chlorophyll maximum (SCM) peaks in a strongly stratified estuary. Could extension and depth location of the SCM be explained by the physical conditions in terms of water column stratification and density interface? Questions were addressed by obtaining data on water column density (CTD), chlorophyll a (Chl a), nutrients, (F _v/F _m), σ_PSII and K _d(PAR) at 15 positions along a 575 km transect in the Kattegat estuary. Results showed that the estuary was strongly stratified with mixed surface and bottom layers intercepted by a layer where density increased with depth. The SCM occurred only in this density interface, and widths of SCM and density interface were highly correlated. The surface waters were nearly depleted of inorganic nitrogen, phosphate and silicate though with significant higher concentrations in the waters below the interface. The Chl a concentration was comparatively higher in the SCM peak as well as maximum quantum efficiency (F _v/F _m) and functional cross sectional area (σ_PSII). The SCM was maintained at very low light levels and by a diapycnal nitrogen flux, with a stratified water column and nutrient depleted surface waters as predecessors. It was concluded that the depth location and vertical extension of the SCM in the estuary were closely linked to the physical structure of the water column in terms of density interface and stratification.     

A method for deriving net primary productivity and component respiratory fluxes from tower-based eddy covariance data: a case study using a 17-year data record from a Douglas-fir chronosequence

Conventional gap‐filling procedures for eddy covariance (EC) data are limited to calculating ecosystem respiration (R_E) and gross ecosystem productivity (P_G) as well as missing values of net ecosystem productivity (F_NEP). We develop additional postprocessing steps that estimate net primary productivity (P_N), autotrophic (R_a), and heterotrophic respiration (R_h). This is based on conservation of mass of carbon (C), Monte Carlo (MC) simulation, and three ratios: C use efficiency (CUE, P_N to P_G), R_a to R_E, and F_NEP to R_E. This procedure, along with the estimation of F_NEP, R_E, and P_G, was applied to a Douglas‐fir dominated chronosequence on Vancouver Island, British Columbia, Canada. The EC data set consists of 17 site years from three sites: initiation (HDF00), pole/sapling (HDF88), and near mature (DF49), with stand ages from 1 to 56 years. Analysis focuses on annual C flux totals and C balance ratios as a function of stand age, assuming a rotation age of 56 years. All six C balance terms generally increased with stand age. Average annual P_N by stand was 213, 750, and 1261 g C m⁻² yr⁻¹ for HDF00, HDF88, and DF49, respectively. The canopy compensation point, the year when the chronosequence switched from a source to a sink of C, occurred at stand age ca. 20 years. HDF00 and HDF88 were strong and moderate sources (F_NEP=−581 and −138 g C m⁻² yr⁻¹), respectively, while DF49 was a moderate sink (F_NEP=294 g C m⁻² yr⁻¹) for C. Differences between sites were greater than interannual variation (IAV) within sites and highlighted the importance of age‐related effects in C cycling. The validity of the approach is discussed using a sensitivity analysis, a comparison with growth and yield estimates from the same chronosequence, and an intercomparison with other chronosequences.     

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.     

PHOTOADAPTATION,GROWTH AND PRODUCTION OF BOTTOM ICE ALGAE IN THE ANTARCTIC1

Biomass, chemical composition, growth rates and the photosynthetic response of natural populations of sea ice algae in McMurdo Sound, Antarctica were followed over most of the spring bloom to examine temporal variability under a relatively constant incident irradiance (ca. 1500–1700 μE · m^-2· s^-1 at solar noon). Collection were restricted to bottom 20 cm of the ice sheet in an area with little or no snow (0–5 cm). At low temperature and irradiance these algae normally exhibited low assimilation numbers (ca. 0.1–0.4 mg C · mg Chl^-1· h^-1). Average growth rates (0.02–0.45 d^-1), based on changes in standing stocks, were also low. Biomass, biochemical composition, growth rates, assimilation numbers and photosynthetic efficiencies (mg C · mg Chl^-1· h^-1 (μE · m^-2· s^-1)^-1) displayed large fluctuations over periods of several days during the growth season. On the other hand, I_k which is an index of photoadaptation, and I_m, the optimal irradiance for photosynthesis, were relatively constant with less than twofold variation throughout our study. Substantial nutrient fluxes (3.3–8.0 mmol Si or N · m^-2· d^-1) were necessary to satisfy the minimum nutrient demand for the observed biomass levels and population growth rates; over the 41 days of our study, integrated nutrient demand represented 69–150 mmol N or Si · m^-2, Only 5–25% of this total demand could be met by all of the nutrients in the ice sheet, if they were readily available. However, adequate amounts were present in the top few meters of the water column. With small nutrient gradients in surface waters below the sea ice, vertical eddy diffusivities on the order of 3.8–9.3 cm²· s^- should supply sufficient nutrients to meet algal demand.     

Dissolved organic matter in Arctic multi-year sea ice during winter: major components and relationship to ice characteristics

Ice cores were collected between 10.03.93 and 15.03.93 along a 200 m profile on a large ice floe in Fram Strait. The ice was typical of Arctic multi-year ice, having a mean thickness along the profile of 2.56 ±0.53 m. It consisted mostly of columnar ice (83%) grown through congelation of seawater at the ice bottom, and the salinity profiles were characterized by a linear increase from 0 psu at the top to values ranging between 3 and 5 psu at depth. Distributions of dissolved organic carbon (DOC) and nitrogen (DON) and major nutrients were compared with ice texture, salinity and chlorophyll a. DOC, DON, dissolved inorganic nitrogen (DIN), NH₄ ⁺ and NO₂ ^– were present in concentrations in excess of that predicted by dilution curves derived from Arctic surface water values. Only NO₃ ^– was depleted, although not exhausted. High DOC and DON values in conjunction with high NH₄ ⁺ levels indicated that a significant proportion of the dissolved organic matter (DOM) was a result of decomposition/grazing of ice algae and/or detritus. The combination of high NH₄ ⁺ and NO₂ ^– points to regeneration of nitrogen compounds. There was no significant correlation between DOC and Chl a in contrast to DON, which had a positively significant correlation with both salinity and Chl a, and the distribution of DOM in the cores might best be described as a combination of both physical and biological processes. There was no correlation between DOC and DON suggesting an uncoupling of DOC and DON dynamics in multi year ice.