CO2 fixation and translocation in benthic marine algae. III. Rates and ecological significance of translocation in Laminaria hyperborea and L. saccharina

Translocation of ¹⁴C-labelled assimilates in Laminaria hyperborea and L. saccharina is strictly basipetal within the thallus. The lowermost 20 cm of the frond accumulates 71% (L. hyperborea) or 63% (L. saccharina) of the basipetallymoving translocate after a period of 96 h from initial feeding with NaH¹⁴CO₃. Whereas translocation was found to occur at all seasons in L. saccharina, which grows throughout the year, no translocation of newly synthesized photosynthate occurred in L. hyperborea in October, when no frond enlargement takes place, nor in January, when the new fond occurs as a tiny outgrowth. In L. hyperborea, organic material, assimilated by the old frond during the period of rapid growth, is exported at the rate of 0.5 mg C · dm^-2 · h^-1. In L. saccharina, 14 to 18 cm frond length are necessary for an elongation rate of the growing zone, situated in lowermost 10 cm of the frond, of 1 cm per week. The significance of translocation in the two Laminaria species is demonstrated by comparison of export rates with growth rates measured in situ and in the laboratory.Dedicated to Professor Dr. Otto Stocker on his 85th birthday.     

The detrital food chain based on seaweeds. I. Bacteria associated with the surface of Laminaria fronds

The bacteria associated with the surface of fronds of the sublittoral brown alga Laminaria longicruris were investigated over a 13-month period on the coast of Nova Scotia (Canada). A psychrophilic population was found to be associated with the frond during the winter and a mesophilic population with the decaying frond during the summer. Numbers of psychrophiles varied inversely with ambient water temperature, and were present in the greatest number on the eroded tips of fronds. Laminaran hydrolyzing isolates were characteristic of the psychrophilic flora and a group of isolates hydrolyzing mannitol, protein and alginate characteristic of the mesophilic population. Increases in the numbers and proportions of bacteria utilizing plant substrates were found to accompany macroscopic evidence of frond decomposition. A comparison of the bacterial floras of L. longicruris fronds from a sheltered and exposed location showed them to be quantitatively and qualitatively comparable.     

Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera in a low-nitrogen environment

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.     

A survey of translocation in laminariales (Phaeophyceae)

A survey of translocation of photoassimilates in 13 genera of Laminariales is presented. All showed long-distance transport of ¹⁴C-labeled products from mature source tissue to meristematic sinks (haptera and intercalary growing regions). In plants with several laminae forming one frond, older laminae may provide assimilates for the growth of younger ones, and in Macrocystis spp., where fronds of different ages and developmental stage arise from a common holdfast, mature fronds initiate and support new fronds. Translocation velocities vary from species to species but are in the range of 55 to 570 mm/h. The results strongly support the hypothesis that Laminariales in general have an effective translocation system, on which their thallus growth depends.     

Growth and DNA damage in young Laminaria sporophytes exposed to ultraviolet radiation: implication for depth zonation of kelps on Helgoland (North Sea)

Growth as an integrative parameter of all physiological processes was measured in young sporophytes of temperate Laminaria digitata, Laminaria saccharina and Laminaria hyperborea exposed in the laboratory to irradiance consisting of either only photosynthetically active radiation (PAR) or to a spectrum including ultraviolet radiation (UVR) (PAR+UVA+UVB) by use of cut-off glass filters. Size increment was measured every 10 min over a period of 18–21 days using growth chambers with on-line video measuring technique. In the chamber, plants were grown at 10±2°C and 16:8 h light–dark cycles with 6 h additional UVR exposure in the middle of the light period. Tissue morphology and absorption spectra were measured in untreated young sporophytes while chlorophyll a content and DNA damage were measured in treated thalli at the end of the experiment. Sensitivity of growth under UVR was found to be related to the observed upper depth distribution limit of the upper sublittoral L. digitata, upper to mid sublittoral L. saccharina and lower sublittoral L. hyperborea. Tissue DNA damage is, however, dependent on thallus thickness which minimizes UVR effect where outer cell layers shade inner cells and provide longer pathlength for UVR. Exposure to UVR causes cellular, enzymatic and molecular damage. Presence of UV-absorbing compounds further reduces effective UVR from reaching physiological targets. The cost of producing higher amount of UV-absorbing compounds and effective DNA repair mechanism can, however, divert photosynthate at the expense of growth. Tissue chlorophyll a content was not significantly different between treatments suggesting a capacity for acclimation to moderate UVR fluence. Growth acclimation to repeated UVR exposure was observed within a period of 12 days while growth inhibition was observed after a longer UVR exposure period of 21 days. The results give further insight into the effects of UVR on the cellular level and show how ecological parameters such as the upper depth distribution limit are dependent on cellular processes.     

An annual carbon budget for the kelp Laminaria longicruris

Oxygen evolution and uptake by whole thalli of the large marine alga Laminaria longicruris de la Pylaie were measured for 24 h, once every 2 weeks for a year, using large chambers to incubate the plants on the sea bed. Diel rates of photosynthesis and respiration were calculated from these measurements and continuous light measurements were used to extrapolate the data between observation dates. The resulting estimates were combined with measurements of growth and carbon content to give an annual carbon budget for a typical mature plant. Annual net assimilation was 6.8 mgC per cm² of frond surface (71 cal cm^-2). Approximately 45% of this appeared in the production of new frond tissue, and a further 12% was accounted for by storage of carbon in mature frond tissue. About 8% was needed for stipe growth, and the remaining 35% was assumed to be lost as dissolved organic carbon. Diel net photosynthetic rates reached a maximum in June and July and were negative only in November, indicating an ability to produce a photosynthetic surplus throughout winter. In early winter the plants drew on stored reserves to supplement photosynthesis in providing carbon for growth, but from January onwards photosynthesis provided more than enough carbon for growth.     

Growth and production of the red alga Gelidium sesquipedale off the Basque coast (northern Spain)

Several aspects of the growth of the agarophyte Gelidium sesquipedale Clemente (Thuret) were studied in sublittoral populations of the Basque coast (northern Spain) from 1981 to 1986. The standing stock of the populations varied greatly with season. During two consecutive years, 60 to 70% of summer biomass had been lost by the end of the unfavourable period of year to growth (autumn and winter). The decline in biomass was mainly due to partial losses of branches, with more branches being lost from larger fronds. Yearly variations in the standing stock were also recorded. Production estimates obtained by a destructive sampling method were not satisfactory because of the spatial heterogeneity of the populations at the three locations studied. Indirect estimation of production by measuring the growth of marked fronds in a control quadrat gave more reliable estimates. Production values of 622±124, 735±147 and 615±123 g dry wt m^-2 yr^-1 were obtained at the three locations, respectively. Frond elongation rates of 7.5, 6.1, 7.5 and 7.9 cm yr^-1, respectively, were recorded during four consecutive years. In winter elongation rates of fronds were rather low. Elongation did not seem to be affected by frond length, the self-shading effect of the canopy, or changes in depth. Cuts resulting from grazing were frequent on fronds through the year, decreasing significantly with increasing depth. Sheared fronds regenerated promptly, independent of season or height at which the cut was made. For commercial purposes the best season for harvesting is summer, which is the period of maximum standing stock, infertility, and calm warm waters that provide easier working conditions for divers.     

Distribution patterns of meiofauna associated with a sublittoral<Emphasis Type="Italic"> Laminaria</Emphasis> bed in the Cantabrian Sea (north-eastern Atlantic)

This study deals with meiofauna associated with a sublittoral population of the kelp Laminaria ochroleuca located on the northern coast of Spain. By sampling once a year over a 4-year period, we examined patterns of faunal distribution as a function of some environmental factors at the meso-scale level (depth, and exposure to waves and surge). We also examined the relationship between L. ochroleuca abundance (as dry weight biomass and number of plants per sampling quadrat) and abundance and diversity of meiofauna. Finally, we investigated patterns of within-plant distribution (algal frond vs. algal holdfast), using also the meiofauna of the adjacent bottom as a referent to estimate the level of "phytal dependence" of the meiofauna collected on L. ochroleuca. We found that the bulk of permanent meiofauna consisted of nematodes, copepods, mites, polychaetes, tanaids and ostracods, with copepods being predominant on the fronds of the alga and nematodes in the holdfasts. The temporary meiofauna consisted of juvenile amphipods, bivalves and gastropods, together with barnacle nauplii and cyprids. Abundance and major composition of meiofaunal taxa were unrelated to both depth and hydrodynamic exposure of the sampling quadrats. However, we detected significant qualitative and quantitative faunal differences as a function of microhabitat. All meiofaunal groups were more abundant in holdfast samples than in frond and bottom samples. The gross taxonomic composition of meiofauna in bottom samples was similar to that in holdfast samples, but substantially different from that of meiofauna associated with the fronds. The L. ochroleuca holdfasts, in which dense aggregations of meiofauna can occur, appear to function as ecotone between phytal and rocky-bottom microhabitats. All together, our results suggest that the distribution of meiofauna within the Laminaria bed is mostly affected by factors operating at the microhabitat level rather than the meso-scale level.Communicated by L. Hagerman, Helsingør     

Demographic variation in the kelp<Emphasis Type="Italic"> Laminaria hyperborea</Emphasis> along a latitudinal gradient

We compared populations of Laminaria hyperborea from different regions along the Norwegian coast (58–71°N). The aim of the study was to examine possible regional differences in population demography and to relate these to latitudinal gradients in ambient temperature and light. A number of population parameters were examined in understory and canopy kelp individuals. We found latitudinal differences in stipe growth rate and growth pattern, mortality, longevity, recruitment and density. Stipe growth rate was higher for young individuals in mid-Norway than in south and north Norway. Mean stipe length was related to temperature and light, and correlated significantly with indices combining temperature during the growth season and daylength in summer. Mortality decreased and longevity increased with increasing latitude, probably because of a temperature decrease with increasing latitude and a general reduction in metabolic rate. Greater longevity may also cause the observed decrease in recruitment rate with increasing latitude, since there are fewer opportunities for understory individuals to replace dead canopy individuals. The development of particularly large kelp in mid-Norway appears to be explained by high growth rates and not a particularly long life span, as earlier assumed. This has consequences for decisions of the optimal harvesting regime for the species in the region, with regard to yield and maintenance of associated biodiversity.Communicated by M. Kühl, Helsingør     

Autecology of Fucus distichus ssp. distichus (Phaeophyceae: Fucales) in Nova Scotia,Canada

The autecology of Fucus distichus L. ssp. distichus was investigated at several sites on the Atlantic coast of Nova Scotia, Canada, throughout a 2-year period. This species is confined to high-littoral rock pools at exposed sites, where there appears to be little competition from other organisms. Water temperature, salinity, dissolved oxygen, and hydrogen-ion concentrations underwent both considerable seasonal and short-term variation. Pools with extremes of salinity did not contain F. distichus ssp. distichus. Receptacles are formed during winter, and by late spring they are cast. Sporelings were first visible during September; their development was slow, and these plants did not become reproductive until the second year. During the period February to May, rapid growth of new fronds occurred; an increase in both length and bulk of the plants resulted from this growth. Fronds of mature plants continued to elongate slowly throughout the summer and the early autumn with the formation of receptacles in winter. Plants were frequently damaged, which resulted in regeneration of new fronds from the wounded surface.Issued as NRCC No. 14510     

Seasonal growth of the giant kelp Macrocystis pyrifera in New Zealand

The seasonal growth pattern of Macrocystis pyrifera (L.) C. Agardh in New Zealand was determined from measurements of blade-elongation rates between March 1986 and December 1987. Growth rates remained relatively constant throughout most of the year, but declined significantly during the summer months. Seawater nitrate levels had a marked seasonal cycle, with concentrations of 0.3 μM detected in summer. The timing of the decline in internal nitrogen concentrations varied for different-aged blades, but occurred ∼1 mo after the decrease in seawater nitrate concentrations. Sufficiently high irradiance levels and seawater nutrient concentrations support active growth for most of the year, but growth is nutrient-limited during summer. No carbon accumulation during summer was observed. Between March and August 1988, growth estimates were also derived from node-initiation rates and stipe-elongation rates to permit comparisons with previous studies from the northern and southern hemispheres. Direct comparisons of the three growth parameters determined for non-terminated canopy and sub-surface fronds were used to assess possible differential resource allocation in the two frond classes. Received: 16 May 1997 / Accepted: 28 May 1997     

Standing crop and leaf area index of the sublittoral Laminaria species near Helgoland

Standing crop values of Laminaria species sampled between 0 to 2 m depth in the sublittoral of Helgoland, a small rocky island in the southern North Sea, are similar to those reported by other workers from Norwegian or Scottish coasts. Below 2 m water depth, however, standing crop values are found to be reduced to a far greater extent with increasing depth than in the Norwegian or Scottish areas studied. Maximum standing crop values are not found in Laminaria digitata and L. saccharina growing just below the low water mark of spring tides, but in the upper L. hyperborea forest at 2 m depth. Population density and leaf area index (LAI) decrease with increasing depth (but not the average weight of individuals constituting the L. hyperborea forest). It is suggested that the depth at which LAI becomes 1 represents an important ecological threshold, since growth rates of L. hyperborea specimens appear to decrease at greater depths but not to increase above the threshold. Self-shading might equalize, to a certain extent, the quantity of underwater irradiance available to an individual phylloid at all depths at which a close vegetation of Laminaria species occurs.     

Morphological variation and phenotypic plasticity of buoyancy in the macroalga Turbinaria ornata across a barrier reef

Many aspects of morphology of benthic algae (length, surface area-to-volume ratio, and blade undulation) are plastic traits that vary in response to physical factors (such as light or water flow environment). This study examines whether frond buoyancy is a plastic trait, and whether differences in morphology including buoyancy affect the potential persistence of macroalgae in habitats characterized by different water flow regimes. Fronds of the tropical alga Turbinaria ornata in protected backreef environments in Moorea, French Polynesia possess pneumatocysts (gas-filled floats) and experience positive buoyant forces, whereas fronds in wave-exposed forereef sites either lack pneumatocysts entirely or have very small, rudimentary pneumatocysts and experience negative buoyant forces. Forereef fronds transplanted to the backreef developed pneumatocysts and experienced increased buoyant force indicating that buoyancy is a phenotypically plastic trait in T. ornata. In comparing the potential for dislodgement by drag, drag was greater on forereef fronds at low flow speeds as these fronds were stiffer and did not bend over at low flow speeds and therefore were less streamlined in the flow than backreef algae, which bent easily. The environmental stress factor (ESF) (a measure of the likelihood of detachment for a frond in its habitat) was higher for forereef than backreef fronds at all flow speeds. When examined with respect to the flow velocities likely in their respective habitats however, the chance of detachment for backreef and forereef was similar. Neither backreef nor forereef fronds were predicted to break under normal, non-storm conditions, but both were predicted to break in storms. Strong forereef morphologies are well suited to habitats characterized by rapid flow, whereas the weaker, buoyant, tall backreef fronds are well suited to habitats where crowding and shading is common but hydrodynamic forces are low.     

Preferences of six leaf beetle species among qualitatively different leaf age classes of three Salicaceous host species

Summary. As Salicaceous plants produce new leaves for a prolonged period of time, they expose a wide range of differentially aged leaves to herbivores during the growing season. In this work, I show that young leaves of three Salicaceous species, Populus tremula L., Salix phylicifolia L. and S. pentandra L., contain more nitrogen than conspecific old leaves. In P. tremula and S. pentandra young leaves also contained more low-molecular weight secondary compounds, phenolic glucosides. Leaves of S. phylicifolia did not contain phenolic glucosides in detectable amounts. Furthermore, in P. tremula and S. pentandra young leaves contained less polymeric digestability-reducing phenolics, condensed tannins, than old leaves. In S. phylicifolia, higher concentrations of condensed tannins were found in young leaves. In laboratory feeding trials with six leaf beetle species, young leaves of the studied plants were invariably preferred in all tested herbivore × host species combinations. In particular, it is remarkable that three leaf beetle species with known different overall relationships to phenolic glucosides equally preferred more glucoside-containing young S. pentandra leaves over conspecific old ones. Four beetle species were found to prefer young leaves of S. phylicifolia despite the higher content of condensed tannins in young leaves. These results indicate that the general preference of leaf beetles for young leaves of Salicaceous plants probably does not primarily result from variable distribution of secondary compounds. Apparently, the preference for young leaves is fundamentally due to variation in leaf nutritive traits, such as nitrogen content. Received 9 February 2001.     

Life history of the sandy-beach amphipod Neohaustorius schmitzi (Crustacea: Haustoriidae)

Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m². Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.     

The annual reproductive cycles of Uca annulipes,Portunus pelagicus and Metapenaeus affinis (Decapoda: Crustacea) from the South-west coast of India

Employing the gonad index method, the reproductive cycles of three decapod crustaceans, Uca annulipes (Latreille), Portunus pelagicus (Linnaeus) and Metapenaeus affinis (Milne-Edwards) have been studied. In these crustaceans breeding is not continuous all the year round, but extends over several months of the year with distinct peak periods of gonadal activity. The male and female reproductive cycles are not concurrent. The peak of the reproductive cycle of males occurs slightly earlier in the breeding season than that of females. These studies indicate the possibility of production of successive broods of eggs during the same breeding season. In these species, the low saline conditions of the monsoon period are unfavourable for breeding. The medium and high saline conditions during the post-monsoon and pre-monsoon months, respectively, with plenty of planktonic food for the larvae, seem to be the favourable periods for breeding activity.     

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.     

Density derived estimates of standing crop and net primary production in the giant kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>

Assemblages of macroalgae are believe to be among the most productive ecosystems in the world, yet difficulties in obtaining direct estimates of biomass and primary production have led to few macroalgal data sets from which the consequences of long-term change can be assessed. We evaluated the validity of using two easily measured population variables (frond density and plant density) to estimate the more difficult to measure variables of standing crop and net primary production (NPP) in the giant kelp Macrocystis pyrifera off southern California. Standing crop was much more strongly correlated to frond density than to plant density. Frond density data collected in summer were particularly useful for estimating annual NPP, explaining nearly 80% of the variation in the NPP from year to year. Data on frond densities also provided a relatively good estimate of seasonal NPP for the season that the data were collected. In contrast, estimates of seasonal and annual NPP derived from plant density data were less reliable. These results indicate that data on frond density collected at the proper time of year can make assessments of NPP by giant kelp more tractable. They also suggest that other easily measured variables that are strongly correlated with standing crop, such as surface canopy area, might serve as similarly useful proxies of NPP.     

Nitrogen supply,tissue composition and frond growth rates for Macrocystis pyrifera off the coast of southern California

Seasonal changes in ambient NO ₃ ^– and NH ₄ ⁺ , tissue composition (N, C, and C/N ratio), and frond growth rates for Macrocystis pyrifera (L.) Agardh were examined. Ambient NO ₃ ^– showed distinct seasonal variations. Frond growth rates were variable, but showed no clear correlation to ambient NO ₃ ^– . The average N content of plant tissue did, however, show the same seasonal variations as ambient NO ₃ ^– . The longitudinal distribution of total tissue N and various components of tissue N along fronds were also analyzed. Several distinct patterns were found: high levels of protein N at growing tips and elevated levels of soluble N in lower parts of the frond. Free amino acids accounted for a major portion of the soluble N, but neither NO ₃ ^– nor NH ₄ ⁺ accumulated in the plant tissue. The longitudinal distribution of N along the fronds is compared to reported variations in C metabolism, and it is concluded that C and N sourcesink relations do not always coincide and bidirectional translocation may occur.     

Seasonal patterns of growth and storage in Laminaria longicruris in relation to differing patterns of availability of nitrogen in the water

The annual cycles of growth and of internal storage of nitrogen and carbon in Laminaria longicruris were compared at three sites in Nova Scotia, Canada. One site had abundant nitrogen all year round, on account of local upwelling, one had abundant nitrogen for only four months and the last was intermediate in this respect. Where nitrogen was abundant year round, growth followed the seasonal pattern of light, and there was little storage of either carbon or inorganic nitrogen. Where nitrogen was only abundant in the sea in winter, growth accelerated at this time and continued throught early summer while internal reserves of nitrogen were utilized. Carbon was stored in summer and used for growth in the fall. At the intermediate site, larger reserves of nitrogen were built up in winter and a high growth rate was maintained through most of the summer. Smaller carbon reserves were accumulated and growth was minimal in winter. Experiments in which plants were transplanted between sites were not conclusive, but suggested that plants at different sites may have genetic differences.