Ecotypic differentiation in light-related traits of the kelp Laminaria saccharina

Light-related traits were compared for Laminaria saccharina Lamour. collected from three habitats in Maine and New York, USA, with different ambient light regimes. Light-level, expressed as a proportion of surface irradiance (I₀), ranged from 0.04 to 0.32 I₀ in the shallow habitat, but rarely exceeded 0.04 I₀ in the deep and turbid habitats. Juvenile sporophytes collected from each habitat in April, 1985, were grown at four acclimation light-levels (0.065, 0.12, 0.26, and 0.54 I₀) in a common-garden, laboratory experiment. Photosynthesis vs irradiance (PI) parameters, light-harvesting characteristics, and rates of carbon-assimilation and growth were determined for each group of plants. The results indicated that ecotypic differentiation had occurred among the three kelp populations. Photosynthetic capacity (P_max) and photosynthetic efficiency () were generally highest for plants from the turbid habitat, lowest for deep plants, and intermediate for shallow plants. These differences were largely attributable to variations in light-harvesting characteristics. The nature and magnitude of photoacclimation responses also differed among populations. Population differences in photosynthetic parameters resulted in different rates of C-assimilation and growth by plants from shallow, deep, and turbid habitats. Predictions of in situ growth rates indicated that the severity of light-limitation and, therefore, the adaptive significance of efficient light-utilization vary among the three populations. It is concluded that ecotypic differentiation in light-related traits is important to the broad environmetal range of L. saccharina.     

Influence of ultraviolet-radiation on chlorophyll fluorescence and growth in different life-history stages of three species of Laminaria (phaeophyta)

Zoospores, gametophytes, young sporophytes and discs cut from mature sporophytes of Laminaria digitata, L. hyperborea and L. saccharina were exposed in the laboratory to UV-radiation, with a spectral composition and irradiance similar to natural sunlight, for periods ranging from 15 min to 8 d, and were then returned to white light. Germination of zoospores and the growth of gametophytes were reduced after exposures to UV longer than 1 h, whereas UV had little effect on the growth of young or mature sporophytes unless exposure continued for more than 48 h. The variable fluorescence (F _v:F_m) of all stages was strongly reduced immediately after short exposures to UV, but recovered almost completely within 24 h. However, exposure of gametophytes to UV for >4 h resulted in little or no recovery of F _v:F_m, whereas >16 h of UV were required to produce this result in young sporophytes, and >48 h in mature sporophytes. Thus, sensitivity to UV-radiation decreased from gametophytes to sporophytes, and with increasing age of sporophytes, but, in gametophytes, growth appeared to be a more sensitive indicator of UV-damage than F _v:F_m after 24 h recovery. The responses to UV of the zoospores and gametophytes of all three species were similar, but both growth and fluorescence measurements suggested that the sporophytes of L. saccharina were more sensitive to UV than those of the other two species.     

Parthenogenesis,apogamy and apospory in Alaria crassifolia (Laminariales)

In cultures of Alaria crassifolia Kjellman, unfertilized eggs developed normally into haploid sporophytes which differentiated into a holdfast, stipe and blade with a midrib. The terminal cells of a male gametophyte grew apogamously into haploid sporophytes with narrow blades which lack the midrib. Further, diploid gametophytes were formed by apospory from vegetative cells of a diploid sporophyte. They were monoecious and their fertilized eggs developed into tetraploid sporophytes. Nuclear phases of the sporophytes and gametophytes concerned were confirmed by cytological observations.This work was supported by Grant No. 38803 from the Ministry of Education of Japan.     

Effects of ocean temperature on the southern range limits of two understory kelps, <Emphasis Type="Italic">Pterygophora californica</Emphasis> and <Emphasis Type="Italic">Eisenia arborea</Emphasis>, at multiple life-stages

Environmental factors have long been shown to influence species distributions, with range limits often resulting from environmental stressors exceeding organism tolerances. However, these abiotic factors may differentially affect species with multiple life-history stages. Between September 2004 and January 2006, the roles of temperature and nutrient availability in explaining the southern distributions of two understory kelps, Pterygophora californica and Eisenia arborea (Phaeophyceae, Laminariales), were investigated along the coast of California, USA and the Baja California Peninsula, Mexico, by limiting either: (a) tissue nitrogen uptake and storage by adult sporophytes during periods of elevated temperature, and/or (b) production of embryonic sporophytes by microscopic gametophytes. Results suggest that while adult sporophytes of both species are tolerant of high temperatures and low nutrients, reproduction by their microscopic stages is not. Specifically, while E. arborea produced embryonic sporophytes at both 12 and 18°C, temperatures commonly observed throughout the southern portion of its range, P. californica produced sporophytes at 12 but not at 18°C. As a result, it appears that the southern distribution of P. californica, which ends in northern Baja California, Mexico, may be limited by temperature acting on its microscopic stages. In contrast, the ability of E. arborea’s microscopic and adult stages to tolerate elevated temperatures allows it to persist in the warmer southern waters of Baja California, as well as to the north along the California coast where both species co-occur.     

Photosynthetic characteristics of giant kelp (Macrocystis pyrifera) determined in situ

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.     

Reproduction,growth and photosynthesis of gametophytes of Laminaria saccharina grown in blue and red light

The induction of reproductive activity by blue light in female gametophytes of the brown alga Lamnaria saccharina has been investigated and related to other effects of red and blue light on these plants. Although germination of zoo-spores is slightly delayed in red light, female gametophytes grow at similar rates in blue and red light of the same quantum irradiance up to an age of 8 to 10 days, when plants in blue light begin to form eggs. The percentage of plants forming eggs is proportional to the total quanta of blue wavelengths received up to a saturating value of about 400 μE·cm^-2; a 50% response is induced by 200 μE·cm^-2. The action spectrum for the induction of fertility has a main peak at 430 to 450 nm, with two subsidiary peaks in the blue and one in the near-ultraviolet light, but the action spectrum for photosynthesis is quite different, with peaks in the red, green and blue regions. These results indicate that egg formation in female gametophytes occurs as a specific morphogenetic response to blue light, which is independent of photosynthesis and growth, and trial experiments with male gametophytes of L. saccharina and gametophytes of other species of Laminariales indicate that these plants react similarly.     

Photosynthetic responses and daily carbon balance ofColpomenia peregrina: Seasonal variations and differences between intertidal and subtidal populations

The seasonal photosynthetic responses and daily carbon gain of upper intertidal, low intertidal and subtidal (3 to 4 m depth) populations ofColpomenia peregrina were examined over a 2 yr period (1986–1988) in Santa Catalina Island, California, USA. The populations showed significant differences in their photosynthetic responses, daily carbon balance and carbon-specific growth rates when normalized to tissue area or to chlorophyll content. The substantial plasticity with respect to photosynthetic responses shown byC. peregrina is considered to be an important factor in facilitating the colonization of both intertidal and subtidal habitats. This species appears to have a cellular carbon metabolism influenced by responses to season and tidal elevation. Highest net daily carbon balance, predicted carbonspecific growth rates and net growth efficiency were achieved in upper intertidal habitats during summer. These parameters decreased in winter and progressively declined with increasing depth as plants become increasingly exposed to low-light regimes. The diminishing net daily carbon balance and predicted carbon-specific field growth rates found during winter suggest that standing stock and lower subtidal limits of distribution ofC. peregrina are at least partly controlled by these two factors.     

Temperature tolerances of two southern African Ecklonia species (Alariaceae: Laminariales) and of hybrids between them

The temperature tolerances in culture of Ecklonia maxima (Osbeck) Papenf., the dominant kelp in the Benguela upwelling region on the west coast of southern Africa, and the smaller E. biruncinata (Bory) Papenf., growing on the warmer south coast, have been investigated. Vegetative growth and reproduction of gametophytes show very similar patterns in the two species, although temperature optima in E. biruncinata are consistently 2 ° to 3 °C higher than those of E. maxima. Reciprocal crosses between clonal female and male gametophytes of the two species produced healthy juvenile sporophytes, indistinguishable from the parent crosses. All parents and hybrids grew well over a wide range of temperatures (8 ° to 22°C), with E. biruncinata sporophytes again having slightly higher optima than E. maxima. Both reciprocal crosses exhibited optima intermediate between the parent crosses. The results are discussed with respect to the origins and evolution of Ecklonia species in the region.     

Influence of local environmental conditions on the seasonal acclimation process and the daily integrated production rates of Laminaria digitata (Phaeophyta) in the English Channel

Two populations of Laminaria digitata (Hudson) Lamouroux (Ann Mus Hist Nat Paris 20:21–47, 1813) were examined for their seasonal photosynthetic acclimation to clear and turbid-light environments along the French coast of the English Channel. Photosynthesis–irradiance curves, pigment concentrations and the daily in situ integrated oxygen production rates were measured in both populations. Despite the great differences in light attenuation between the sites, the two populations achieved similar oxygen production rates in the field, in relation to high maximal photosynthetic rates, total pigment concentrations and antenna (fucoxanthin + chlorophyll c)/chl a pigment ratios in sporophytes from the turbid environment. Environmental conditions (i.e. light, temperature and nitrogen availability) changed throughout the year in both sites. While the seasonal acclimation trends were evident in the clear-light environment, the strategy in the turbid-light environment differed, tending to maximize light capture throughout the year. This study highlights the diversity of the response of a single species to contrasted light environments.     

Effect of blue light on early sporophyte development of Saccharina japonica (Phaeophyta)

Growth and photosynthesis in the blade of Saccharina sporophytes are strongly stimulated by blue light. However, little is known about the effect of blue light on the early development and longitudinal photosynthesis profile of Saccharina sporophytes. In this study, S. japonica sporelings were cultured under blue or red light for 8 weeks. Blue light affected longitudinal, tangential cell divisions and tissue differentiation early in sporophyte development. The number of latitudinal cells in the blade under blue light was over fivefold that under red light. In addition, the number of cell layers was higher in the growing point than in the blade under blue light, whereas sporelings grown under red light contained only a single cell layer. Under blue light, the photosynthetic capacities of the growing region, blade and stipe were similar, and the maximum relative electron transport rate was even lower in the growing point than in the blade. The longitudinal photosynthesis profile suggested that blue light stimulated the enzymes participating in light-independent carbon fixation in the growing point and accordingly was less dependent on high light irradiances. Collectively, the results indicated that blue light promotes the early development of S. japonica sporophytes, which was attributed to both photomorphogenetic responses and photosynthetic reactions.     

Size and spatial structure in deep versus shallow populations of the Mediterranean gorgonian <Emphasis Type="Italic">Eunicella singularis</Emphasis> (Cap de Creus,northwestern Mediterranean Sea)

In the Western Mediterranean Sea, the gorgonian Eunicella singularis (Esper, 1794) is found at high densities on sublittoral bottoms at depths from 10 to 70 m. Shallow colonies have symbiotic zooxanthellae that deeper colonies lack. While knowledge of the ecology of the shallow populations has increased during the last decades, there is almost no information on the ecology of the deep sublittoral populations. In October and November 2004 at Cap de Creus (42°19′12″ N; 03°19′34″ E), an analysis of video transects made by a remotely operated vehicle showed that shallow populations (10–25 m depth) were dominated by small, non-reproductive colonies, while deep sublittoral populations (50–67 m depth) were dominated by medium-sized colonies. Average and maximum colony heights were greater in the deeper populations, with these deeper populations also forming larger patch sizes and more extensive regions of continuous substrate coverage. These results suggest that shallow habitats are suitable for E. singularis, as shown by the high recruitment rate, but perturbations may limit or delay the development of these populations into a mature stage. This contrasts with the deep sublittoral habitats where higher environmental stability may allow the development of mature populations dominated by larger, sexually mature colonies.     

Photoadaptation of zooxanthellae in the sponge Cliona vastifica from the Red Sea, as measured in situ

In the Red Sea, the zooxanthellate sponge Cliona vastifica (Hancock) is mainly present at >15 m depth or in shaded areas. To test whether its scarcity in unshaded areas of shallower waters is linked to the functional inefficiency of its photosymbionts at high irradiances, sponges were transferred from 30 m to a six times higher light regime at 12 m depth, and then returned to their original location. During this time, photosynthetic responses to irradiance were measured as rapid light curves (RLCs) in situ by pulse amplitude modulated (PAM) fluorometry using a portable underwater device, and samples were taken for microscopic determinations of zooxanthellar abundance. The zooxanthellae harboured by this sponge adapted to the higher irradiance at 12 m by increasing both their light saturation points and relative photosynthetic electron transport rates (ETRs). The ETRs at light saturation increased almost fourfold within 15–20 days of transfer to the shallower water, and decreased back to almost their original values after the sponges were returned to 30 m depth. This, as well as the fact that the photosynthetic light responses within an individual sponge were in accordance with the irradiance incident to specific surfaces, shows that these photosymbionts are highly adaptable to various irradiances. There was no significant change in the number of zooxanthellae per sponge area throughout these experiments, and the different photosynthetic responses were likely due to adaptations of the photosynthetic apparatus within each zooxanthella. In conclusion, it seems that parameters other than the hypothesised inability of the photosymbionts to adapt adequately to high light conditions are the cause of C. vastifica's rareness in unshaded shallow areas of the Red Sea. Received: 25 April 2000 / Accepted: 13 October 2000     

Effects of water motion on propagule release from algae with complex life histories

Reproductive marine algae with complex life histories may respond differently to water motion depending upon whether the spore-producing or gamete-producing phase is considered. Two such species, the kelp Alaria esculenta (L.) Greville (Phaeophyceae) and Ulva lactuca L. (Chlorophyta), were examined experimentally in the laboratory. The kelp was collected in April–June 2000 and U. lactuca was collected in July–August 2000, from four intertidal habitats at Schoodic Point, Maine, USA. Orbital shakers were used to generate water motion. Sporophylls of A. esculenta released more zoospores under shaken versus calm conditions, whereas fewer antheridia on the microscopic male gametophytes released sperm under shaken versus calm conditions when male and female gametophytes were placed together. However, antheridial sperm release was equivalent when male gametophytes were exposed to undiluted media from dense cultures of mature female gametophytes under shaken versus calm conditions. These data suggest that water motion inhibited sperm release by diluting the sperm-releasing pheromone produced by ripe eggs below a threshold required to cause sperm release. Water motion stimulated both gamete and zoospore release from U. lactuca. This is the first report in an alga of stimulation of gamete release by increased water motion, but it is notable that parthenogenesis occurs in Ulva spp.; thus, gametes may develop into gametophytes (1 N) or parthenosporophytes (2 N). This study demonstrates that water motion has strong effects upon algal reproduction, but that the effects may vary between species, possibly due to their different life histories.Communicated by J.P. Grassle, New Brunswick     

Polymorphism in the brown alga<Emphasis Type="Italic"> Dictyota dichotoma</Emphasis> (Dictyotales,Phaeophyceae) from Korea

Two morphotypes of Dictyota dichotoma of uncertain taxonomic status have been identified in Korea. The broad thallus type is common on the south and east coasts of Korea, and is similar to European D. dichotoma in gross morphology. The slender thallus type is only found on the west coast. Morphological differences between the two plant types are evident in their thallus width, their terminal bifurcations, the shapes of apices, and their thallus thickness. The differences in the size and shape of the tetrasporangia and tetraspores are also notable. Plants from the east coast exhibit a dominant asexual annual life-cycle involving sporophytes but not gametophytes, and maintain their population by the production of in-situ germlings. In the slender plants from the west coast, a sexual life-cycle dominates. The rbcL, rbcS, psaA and psbA gene sequence differences between the two morphotypes were congruent with morphological, anatomical and phenological characteristics. In spite of these differences, there is no known inbreeding barrier between these two morphotypes. The L_m/L_c values of the second internodes from two types are also within the range of the species criterion specified by Hörnig et al. These results suggest that the two types of D. dichotoma from Korea should be classified as a single species and that they may be in the process of speciation due to geographical isolation (allopatric speciation).Communicated by O. Kinne, Oldendorf/Luhe     

Changes in morphology and physiology of an East Mediterranean sponge in different habitats

The sponge Tetilla sp. (Tetractinomorpha: Tetillidae) is a common species in the eastern Mediterranean. This sponge inhabits four different habitat types differing in wave impact and irradiance levels. Two of these habitats (a shallow cave and deep water) are characterized by relatively calm water, whereas the other two (shallow exposed site and tide pools) are in turbulent water with high energy flow. The present study examined the influence of physical (depth, illumination and water motion) and biotic factors on morphology, skeletal plasticity and reproductive traits among the four spatially separated populations. Sponges from tidal pools had significantly larger body volume than sponges from deep water and from shallow caves (ANOVA: tidal-deep P<0.0001; tidal-shallow caves P<0.05). Sponges from exposed habitats were significantly larger than deep-water sponges (ANOVA: P=0.01). In addition, individuals from tide pools and from the exposed habitat had a significantly higher proportion of structural silica than sponges from the calmer deep water and from the cave sites. Oxea spicules in sponges from the calm habitats were significantly shorter than in those from the tidal pools and the exposed habitats. The percentage of spicules out of a sponges dry weight in individuals transplanted from deep (calm) to shallow (turbulent) water significantly increased by 21.9±12.9%. The new spicule percentage did not differ significantly from that of sponges originally from shallow water. Oocyte diameter differed significantly between habitats. The maximal size of mature eggs was found in deep-water sponges in June (97±5 m). In the shallow habitats, a smaller maximal oocyte diameter was found in the cave, in May (56.5±3 m). Furthermore, oocyte density in shallow-water sponges was highest in May and decreased in June (with 88.2±9 and 19.3±9 oocytes mm^–2, respectively). At the same time (June), oocyte density of deep-water sponges had just reached its maximum (155±33.7 oocytes mm^–2). The difference in oocyte size and density between deep- and shallow-water individuals indicates an earlier gamete release in the shallow sponge population. The results suggest that plasticity in skeletal design of this sponge indicates a trade off between spicule production and investment in reproduction.Communicated by O. Kinne, Oldendorf/Luhe     

Temperature ecotypes near the southern boundary of the kelp Laminaria saccharina

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.     

Photoadaption in marine phytoplankton: Response of the photosynthetic unit

Some species of phytoplankton adapt to low light intensities by increasing the size of the photosynthetic unit (PSU), which is the ratio of light-harvesting pigments to P700 (reaction-center chlorophyll of Photosystem I). PSU size was determined for 7 species of marine phytoplankton grown at 2 light intensities: high (300 E m^-2 s^-1) and low (4 E m^-2 s^-1); PSU size was also determined for 3 species grown at only high light intensity. PSU size varied among species grown at high light from 380 for Dunaliella euchlora to 915 for Chaetoceros danicus. For most species grown at low light intensity, PSU size increased, while the percentage increase varied among species from 13 to 130%. No change in PSU size was observed for D. euchlora. Photosynthetic efficiency per chlorophyll a (determined from the initial slope of a curve relating photosynthetic rate to light intensity) varied inversely with PSU size. In contrast, photosynthetic efficiency per P700 was enhanced at larger PSU sizes. Therefore, phytoplankton species with intrinsically large PSU sizes probably respond more readily to the rapid fluctuations in light intensity that such organisms experience in the mixed layer.Contribution No. 1180 from the Department of Oceanography, University of Washington, Seattle, Washington, USA     

Optimal growth and maximal survival temperatures of Atlantic Laminaria species (Phaeophyta) in culture

The effects of temperature on growth rate of rapidly-growing cultured macrosporophytes of 9 isolates of Atlantic Laminaria comprising 4 species have been investigated. No significant population variation was observed within species despite wide variations in temperature between the original collecting sites. L. saccharina showed a broad temperature optimum in the 10°–15°C range, whereas L. longicruris had a sharp optimum at 10°C. L. digitata and L. hyperborea grew more slowly, with only slightly sub-optimal growth over a wide temperature range, but with peaks at 10°C (L. digitata) and 15°C (L. hyperborea). The maximum survival temperatures of individual male and female vegetatively-growing gametophytes were ascertained for these species plus the Arctic L. solidungula, and were as follows: L. saccharina and L. longicruris, 23°C; L. digitata (male), 23°C; L. digitata (female), 22°C; L. hyperborea, 21°C; L. solidungula, 18°C. The lack of within-species differences demonstrates that the success of the genus in areas with different temperature regimes is brought about by phenotypic plasticity of individuals rather than the selection of temperature races or ecotypes.     

Effects of high-frequency light fluctuations on growth and photoacclimation of the red algaChondrus crispus

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.