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Notes: Abstract Effects of high irradiance on photosynthetic characteristics were examined in sporophytes of the kelp Laminaria saccharina Lamour. from 1992 to 1994. Exposure to high irradiance (700 μmol photons m-2s-1) for 1 h at optimal temperature (12°C) caused a 40 to 60% decline in photosynthetic efficiency (alpha), quantum yield, and the ratio of variable to maximum chlorophyll fluorescence (Fv/Fm), an indicator of Photosystem II efficiency. Although the photoinhibition effects were partly attributable to protective mechanisms, a concurrent increase in minimal fluorescence (Fo) indicated damage to Photosystem II reaction centers. The magnitude of photoinhibition was proportional to irradiance and duration; however, Fv/Fm was significantly reduced after exposure to irradiances as low as 40 to 50 μmol photons m-2s-1 for 1 h, or to 700 μmol photons m-2s-1 for only 5 min. In contrast, photosynthetic capacity (Pmax) was affected only at much higher irradiance. Superoptimal temperatures up to 24°C did not exacerbate high-light effects. At 25°C, however, alpha and Pmax were more susceptible to photoinhibition than at lower temperatures. Recovery from photoinhibition was examined by following Fv/Fm and Fo for 24 h after exposure to high light. Recovery of Fv/Fm was fastest during the first 1 to 3 h, and slowed or ceased after 6 to 12 h, while recovery of Fo was relatively constant over 12 h. Dithiothreitol, which blocks formation of energy-dissipating xanthophylls, reduced both the initial rate and extent of recovery. Chloramphenicol, which blocks chloroplast-encoded protein synthesis, had little effect on initial rates of recovery, but stopped recovery after 3 h. Thus, L. saccharina appears to rely on the xanthophyll cycle to protect the photosynthetic apparatus, and reversal of this protective mechanism causes the rapid initial recovery in Fv/Fm. Longterm recovery depends on repair of damaged reaction centers. Both the rate and extent of recovery were temperature-dependent. The initial rate was higher at 18 to 22°C than at 12°C, but the extent of recovery over 24 h declined with increasing temperature. High temperatures, therefore, appear to enhance protective mechanisms, but disrupt repair processes. L. saccharina from Long Island Sound, an ecotype adapted to low light and high temperature, showed slightly but consistently greater effects of photoinhibition than plants from the Atlantic coast of Maine, but exhibited faster recovery at superoptimal temperatures.

Notes: Abstract Rates of net photosynthesis and nocturnal respiration by individual blades of the giant kelp Macrocystis pyrifera (L.) C. Agardh in southern California, were determined in situ by measuring oxygen production in polyethylene bags during spring/summer of 1983. Mature blades from different depths in the water column exhibited different photosynthetic characteristics. Blades from the surface canopy (0 to 1 m depth) exhibited higher photosynthetic capacity under saturating irradiance and higher photosynthetic efficiency at low irradiances than blades from 3 to 5 or 7 to 9 m depths. Saturating irradiance was lower for canopy blades than for deeper blades. Canopy blades showed no short-term photoinhibition, but photosynthetic rates of deeper blades were significantly reduced during 1 to 2 h incubations at high irradiances. Results of 1 to 2 wk acclimation experiments indicated that differences between photosynthetic characteristics of blades from different depths were primarily attributable to acclimation light conditions. Vertical displacement of blades within the kelp canopy occurred on a time-scale of 1 min to 1 h. Blades continually moved between the unshaded surface layer and deeper, shaded layers. Vertical movement did not maximize photosynthesis by individual blades; only a small proportion of blades making up a dense surface canopy maintained light-saturated photosynthetic rates during midday incubations. The relatively high photosynthetic rates exhibited by canopy blades over the entire range of light conditions probably resulted from acclimation to intermittent high and low irradiances, a consequence of vertical displacement. Vertical displacement also reduced the afternoon depression in photosynthesis of individual canopy blades. The overall effect of vertical displacement was optimization of total net photosynthesis by the kelp canopy and, therefore, optimization of whole-plant production.

Notes: Abstract Effects of temperature on survival, growth, and photosynthesis were compared for two USA populations of Laminaria saccharina Lamour. One population was located in New York State, near the southern latitudinal boundary of the species in the western North Atlantic. This southern boundary population was exposed to ambient temperatures ≧20°C for about 6 wk each summer. The second population was located in Maine, toward the center of the latitudinal range of the species, and was rarely exposed to temperatures〉17°C. sporophytes from the New York (NY) population exhibited greater tolerance of high temperature than plants from the Maine (ME) site. Juvenile sporophytes from the two sites had similar rates of survivorship and growth at temperatures below 20°C, but showed different responses at 20°C in laboratory experiments. NY plants survived and grew for 6 wk at 20°C. ME plants showed negative growth during wk 2 and 100% mortality during wk 3. NY and ME plants held in situ at the NY site during June to September, 1985, also exhibited differential survivorship when ambient temperatures exceeded 20°C. Results of photosynthesis and dark respiration measurements on NY and ME plants grown at various temperatures suggested that the high-temperature tolerance of NY plants was attributable to their ability to maintain positive daily net C-fixation at 20°C. The high-temperature tolerance of the NY plants appeared to be due to genetic adaptation and is probably crucial to the persistence of the species near its southern boundary.

Notes: Abstract An adult giant kelp plant (Macrocystis pyrifera), moved from an inshore kelp forest to an offshore, low-nitrogen environment near Santa Catalina Island, California (USA), maintained growth for 2 wk on internal nitrogen reserves. Frond elongation rates decreased significantly during the third week, and plant growth rate (wet wt) dropped from an initial inshore rate of 3.6 to 0.9% d-1. During this 3 wk period, nitrogen contents and free amino acid concentrations decreased, while mannitol and dry contents increased in frond tissues. After depletion of internal nitrogen reserves, the nitrogen content of lamina and stipe tissues averaged 1.1 and 0.7% dry wt, respectively. The experimental plant was exposed to higher ambient nitrogen concentrations during the last 2 wk. Rates of frond elongation and plant growth increased, but nitrogen content and amino acids in frond tissues remained low. Of the total nitrogen contained in frond tissue of the plant before transplantation, 58% was used to support growth in the absence of significant external nitrogen supply. Amino acids constituted a small proportion of these internal nitrogen reserves. Net movement of nitrogen occurred within large fronds, but not between different frond size classes. The nitrogen content of holdfast tissue remained relatively constant at 2.4% dry wt and accounted for 18 to 29% of the total nitrogen. Holdfast nitrogen was not used to support growth of nitrogen-depleted fronds. In comparison to Laminaria longicruris, which is adapted to long seasonal periods of low nitrogen availability, M. pyrifera has small nitrogen-storage capacity. However, internal reserves of M. pyrifera appear adequate to make nitrogen starvation uncommon in southern California kelp forests.

Notes: Abstract Rates of NO 3 - uptake by individual blades of Macrocystis pyrifera (L.) C. Agardh were measured at different flow rates in the laboratory. Dissolution rates of hemispherical, plaster buttons attached to the blade surface provided a relative measure of flow rates over blades used in uptake experiments and also over intact blades of adult kelp plants in situ (Laguna Beach, California, USA; 1981). Laboratory results indicated that uptake was saturated at a flow rate equivalent to 2.5 cm s-1 current velocity. Flow rates over intact blades in situ always exceeded this uptake saturation level. Wave surge and movement of plant surfaces relative to the surrounding water provided sufficient flow to saturate uptake, even in a dense kelp canopy during low-current and calm sea-state conditions.

Notes: Abstract We have developed a population model for the giant kelpMacrocystis pyrifera (L.) C. Agardh in southern California, USA. The model includes five life-history stages and takes into account environmental and demographic stochasticity, as well as density-dependent interactions. The density of each stage is predicted on a monthly basis for up to 20 yr, and extinction probability is determined for adult sporophytes. Survival probabilities and rates of reproduction and growth are based on stage-specific responses to environmental conditions (irradiance and temperature), including the occurrence of El Niño events. The model is validated by comparing simulation results to empirical data from natural kelp populations. Results of the model provide insight into patterns observed in natural populations and have applications in resource management.

Notes: Abstract High-frequency fluctuations due to wavefocusing are prominent characteristics of light in shallow marine environments. Effects of high-frequency (0.01 to 1 Hz) fluctuating light on growth rates of the red algaChondrus crispus Stackh., collected from Crane Neck Point, New York, USA, during July 1988, were determined by comparing plants grown under fluctuating and constant light regimes with similar daily irradiances. At high daily irradiance, growth rates were higher under fluctuating light than under constant light. Fluctuating light effects were frequency-dependent; growth was enhanced by fluctuations at 0.1 and 1 Hz, but not by fluctuations at 0.01 Hz. At low daily irradiance, growth rates were not affected by fluctuating light at any frequency tested. Enhancement of growth was not due to effects of high-frequency light fluctuations on photoacclimation responses ofC. crispus. Plants grown under fluctuating light at high daily irradiance actually exhibited lower photosynthetic capacity and efficiency (determined under constant light) than plants grown under constant light. These differences were attributable to variation in the density of Photosystem II reaction centers, which was low in plants grown under fluctuating light. Maximum turnover rate of whole-chain electron transport and activity of ribulose-1,5-bisphosphate carboxylase were affected by total daily irradiance, but not by high-frequency light fluctuations. Enhancement of growth under fluctuating light was partly attributable to reduced rates of dark respiration compared to rates of plants grown under constant light. The results also provided indirect evidence that high-frequency light fluctuations may enhance instantaneous photosynthetic rates. This effect could increase daily carbon gain and, therefore, stimulate growth ofC. crispus under high-frequency light fluctuations.

Notes: Abstract N2-fixation associated with the green macroalgaCodium fragile subsp.tomentosoides (van Goor) Silva from Long Island, New York, USA, was attributable to several species of endophytic cyanobacteria. Rates of N2-fixation ranged from 0.03 to 3.2µg N g−1 dry wt h−1 in freshly collected plants from several sites. Growth of the cyanobacteria appeared to be light-limited, due to the transmission of only 5 to 10% of incident light through the pigmented surface-layer of the macroalga. Daily irradiance was the most important factor determining both abundance of cyanobacterial cells and rate of N2-fixation. The rate was also affected by instantaneous irradiance, and increased twofold from dark to ambient surface irradiance. Rates were reduced at low temperature (8°C) but showed no temperature effect between 12° and 26°C. External concentrations of dissolved inorganic nitrogen (DIN) up to 20µM did not influence N2-fixation rate, but long-term exposure to 60µmol l−1 d−1 of NH 4 + caused a reduction in the rate. InC. fragile grown under high daily irradiance and low external DIN concentration, ~50% of the assimilated-N was attributable to N2-fixation. However, chlorophyllb extracted from plants grown with15N2 showed an atom % excess15N of less than 0.1, suggesting that only a small proportion of the bacterially fixed-N was transferred to the seaweed. The association betweenC. fragile and its endophytic cyanobacteria appears to be based primarily on microhabitat suitability, rather than mutual metabolic dependence. It is doubtful that N2-fixation by cyanobacteria is important to the ecological success of this seaweed species.

Notes: Abstract Comparison of cultured sporophytes and gametophytes in common-garden experiments confirmed the existence of ecotypic differences in light-related traits among populations ofLaminaria saccharina (L.) Lamour. Cultured sporophytes from the turbid habitat in Long Island Sound, New York, USA, grew faster under both limiting and saturating daily irradiances than sporophytes from shallow and deep habitats along the Atlantic coast of Maine. Rapid growth of turbid plants was attributable to several factors, including high photosynthetic capacity and efficiency [due to differences in photosynthetic unit (PSU) number and size], low respiration rates, and high surface area:weight ratios. In contrast to sporophytes, microscopic gametophytes from the three kelp populations grew at similar rates under limiting and saturating daily irradiances. Biomass-specific photosynthesis vs irradiance (PI) parameters were similar for gametophytes from the shallow, deep, and turbid sites, despite population differences in chlorophyll-specific PI parameters and PSU characteristics. However, turbid gametophytes produced microscopic sporophytes more rapidly than gametophytes from the shallow and deep sites, apparently due to a lower blue-light requirement for gametogenesis. Ecotypic differences in sporophytes and gametophytes ofL. saccharina from shallow, deep, and turbid habitats can be understood as phase-specific adaptations.

Notes: Abstract Quantum irradiance was measured at several depths in forests of the giant kelp Macrocystis pyrifera (L.) C. Agardh in southern California, USA, during the summer of 1983. The greatest reduction in irradiance occurred in the uppermost 1 m of the water column, where the floating portions of kelp fronds formed surface canopies with high blade densities. Average irradiances low enough to limit macroalgal photosynthesis (〈200 μE m-2 s-1) were recorded at 1 m depth below dense kelp canopies under sunny surface conditions. Light penetration was exponentially related to canopy density, but was higher than predicted from transmission through individual kelp blades dae to the heterogeneous distribution of canopy tissue. Light penetration was considerably lower immediately adjacent to an individual kelp plant than in the interplant spaces, indicating significant self-shading. Spatial and temporal variability in both irradiance and light penetration were highest just below the canopy and decreased with increasing depth. Sunflecks (〉200 μE m-2 s-1), ranging from 〈1 to 60s or more in duration, composed a considerable proportion of total irradiance at 1 m depth, even during recordings with low average irradiances. These fluctuations probably affect photosynthetic efficiency of kelp tissues within and just below the canopy. Predictions of kelp productivity based on irradiances averaged over hours or days and on photosynthetic parameters determined under laboratory conditions may be inaccurate, due to the spatial and temporal variability characteristic of the kelp forest light-environment.