Effect of light on apical length growth of Fucus distichus spp. edentatus (Phaeophyta)

The length growth of Fucus distichus L. ssp. edentatus (Pyl.) Powell in continuous light is compared to growth in natural daylight. In continuous light the length growth of vegetative and fertile apices were saturated at 25 to 40 Wm^-2 and 15 to 30 Wm^-2, respectively. In natural daylight vegetative apices reached saturation at about 300 Wm^-2 (in May), while fertile apices were not saturated at 180 Wm^-2 (in March). This shows that saturation levels are dependent on the light regime and that they may be greatly underestimated when continuous light is used. At 20 Wm^-2 vegetative and fertile apices were daylength saturated at 8 and 12 to 16 h, respectively. Maximum growth in vegetative apices occurred in July, but growth rates were high also during wintertime (45% of July maximum in March at 4.0°–4.2°C). This corresponds to the experimental results which indicate that F. d. edentatus grows well at low irradiances and during short photoperiods.     

Nitrogen uptake and growth of the germlings and mature thalli of Fucus distichus

Fucus distichus L. was collected near Vancouver, Canada, in late fall and early winter, 1981. The effects of the forms of nitrogen (nitrate, ammonium or urea) and periodic exposure to air on growth, rhizoid development and nitrogen uptake in germlings was investigated. Gamete release, fertilization, germination and germling growth had no requirement for a specific form of nitrogen. Periodic exposure to air increased secondary rhizoid development twofold. Nitrate and ammonium uptake rates of the germlings were higher than for the mature thalli (20 to 40 times for nitrate and 8 times for ammonium), while the halfsaturation constant (K _s) values for nitrate were similar (1 to 5 M). The germlings showed saturable uptake kinetics but the mature thalli did not. When germlings were exposed to air it caused a 70% decrease in nitrate uptake, but not change in ammonium uptake. Ammonium uptake in the mature thalli was proportional to the ambient ammonium concentration. Nitrate uptake in the mature thalli appeared to follow saturation kinetics at low nitrate concentrations, but showed a non-saturable component at concentrations greater than 10 M. Presence of ammonium inhibited nitrate uptake by the mature plants but not by the germlings.     

Population biology ofMacrocystis integrifolia from British Columbia,Canada

Aspects of the population dynamics ofMacrocystis integrifolia Bory were determined from harvested wild populations over a 26-month period and cultivated plants over a 40-month period. Mean wild plant size varied from 0.1 wet kg and two fronds in the winter to 3.1 wet kg and mine fronds in the summer. Plant recruitment probably occurred year-round with recruits making up 20 to 86% of the total population. Plant half-lives, estimated from population data, ranged from 1 to 3 months, with few plants exceedings 2 to 3 years in age. Standing crop ranged from 1.6 plants m^-2 and 0.1 wet kg m^-2 in winter to 8.3 plants m^-2 and 8 wet kg m^-2 in summer.     

Introduced versus native subspecies of Codium fragile: how distinctive is the invasive subspecies tomentosoides?

After its introduction, the green alga Codium fragile (Sur.) Hariot ssp. tomentosoides (van Goor) Silva has spread widely on several temperate-zone, rocky shores where non-weedy conspecific subspecies occur (N.E. Atlantic, N.E. Pacific, S. Pacific). To determine how phenologically and morphologically distinctive the invasive alga was relative to native subspecies, I compared marine intertidal populations of C. fragile ssp. tomentosoides and the native C. fragile ssp. novaezelandiae (J. Ag.) Silva (hereafter referred to as ssp. tomentosoides and ssp. novae-zelandiae respectively on New Zealand shores in 1992, 1993 and 1995. On the North Island, the invasive ssp. tomentosoides is sparsely distributed on low intertidal benches on wave-protected shores in the Hauraki Gulf (east coast) in spring and summer, and thalli die back to the perennial holdfast in autumn. In contrast, the native ssp. novaezelandiae forms dense beds within the low intertidal mussel zone on wave-swept shores of Maori Bay (west coast), and fronds are perennial. Whereas ssp. tomentosoides has only a few fronds arising from the spongy basal hold-fast, ssp. novae-zelandiae thalli are composed of many fronds. The ssp. tomentosoides from the Hauraki Gulf is significantly more branched than comparably sized native conspecifics from Maori Bay. These phenological and morphological differences were used to predict the subspecific identity of C. fragile from three other locations on the North Island, two locations on the South Island, and four locations on S.E. Australian shores; microscopic examination of utricles was used to check the predictions. Seasonality and number of fronds per thallus are the most reliable characters for field identification of native vs invasive subspecies: perennial intertidal thalli with large numbers of fronds are indicative of native subspecies for different geographic regions.     

The effect of water movement,temperature and salinity on abundance and reproductive patterns of<Emphasis Type="Italic"> Macrocystis</Emphasis> spp. (Phaeophyta) at different latitudes in Chile

This study describes the density variation and phenology of Macrocystis integrifolia and M. pyrifera populations from northern and southern Chile, respectively. Samples of both species were taken in wave-exposed and wave-protected areas. In addition, spore production, germination and early growth rate of sporophytes of each population was studied at monthly intervals under three temperature and salinity regimes. Results indicate that M. integrifolia from northern Chile presents perennial plants with a mean density of three individuals per 0.25 m² throughout the year and that it reproduces mainly during spring and winter. Although, M. pyrifera in exposed areas of southern Chile also have a perennial-type life strategy, they are able to reproduce all year round. In contrast, M. pyrifera populations in protected areas of southern Chile show a clear annual cycle, with high recruitment during late winter and fertile sporophytes in summer and autumn, although the populations become completely decimated thereafter. The effect of temperature and salinity on M. integrifolia shows that it is independent of water movement, but requires low temperatures and high salinities for the release of zoospores, germination and early sporophyte growth. This pattern differs from that of M. pyrifera in southern Chile, which has a broader tolerance range for salinity and temperature than does M. integrifolia. However, in southern Chile wave-protected populations showed higher spore release and germination at 15°C and 18°C, whereas sporophyte growth responded better at the lowest temperature tested (8°C). In general, these results are contrary to those expected, since a seasonal reproductive pattern was observed in M. integrifolia inhabiting a less seasonally variable environment. In exposed sites of southern Chile, plants showed greater tolerance and continuous reproduction throughout the year, despite the greater environmental variability. Finally, population dynamics of protected kelps in southern Chile shows an annual pattern, which is contrary to the expected perennial strategy shown by exposed populations.Communicated by P.W. Sammarco, Chauvin     

Growth of amputated and dark-exposed individuals of the brown alga Laminaria hyperborea

The seasonal growth of the brown alga Laminaria hyperborea (Gunn.) Foslie, which forms dense forests in the sublittoral zone of Helgoland, a rocky island in the Southern North Sea, was investigated by transplanting specimens of medium size onto PVC or wooden plates fixed to sub-tidal grwoth stations. The 2-year study revealed that, during the season of fast growth (January to June), young plants produce each year a new frond, larger in size than in the preceding year. This observation can be interpreted, in part, by assuming that the growth of the young frond is accomplished not only by its own assimilatory surplus, but also by reserve materials of the old frond assimilated during the preceding season of slow growth. This assumption is supported by experimental evidence: (1) Even in complete darkness Laminaria hyperborea is capable of producing a small new frond during the season of fast growth; (2) New fronds with stipe, but without old frond, grow (during the same period) considerably faster than isolated new fronds separated from the possible sources of reserve materials. Maximum growth occurs in normal plants which possess their old frond until April or May.An abbreviated version of this paper has been presented at the Sixth International Seaweed Symposium, held at Santiago de Compostela, Spain, September 9 to 13, 1968.     

Intraspecific geographic-morphological variation patterns in Fucus distichus and F. evanescens

The intraspecific patterns of morphometric variation were examined in Fucus distichus L. and F. evanescens C.Ag., using samples gathered from 49 sites spread across the North Atlantic. Ten characters were measured on each plant. The data were subjected to bivariate regression analyses, canonical correlation analysis and nested analysis of variance (ANOVA). It was shown that the combined effects of the three environmental variables measured for each site (latitude, longitude and exposure) could only explain 23 and 15% respectively of the intraspecific morphometric variation. Inter-individual, within-site differences accounted for a further 62 and 53%. The remaining variation appears to be expressed on a geographic scale of tens of kilometres at most. Thus, the species seem to take the form of fine-scale phenotypic mosaics, superimposed on weak major clines. Such a pattern has not previously been described for any macroalga, although it is known in some angiosperms.     

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.     

Seasonal growth and composition of fronds of Macrocystis pyrifera in the Falkland Islands

Frond growth of Macrocystis pyrifera in the Falkland Islands was monitored in shallow coastal water from December 1985 to March 1987, and at a different site in deeper water from December 1985 to June 1986. Growth rates in the deeper bed were generally higher than those recorded in the coastal zone. At both sites, node initiation and elongation rate fluctuated according to the seasonal pattern of light or water temperature. In the shallow coastal area, nitrate was abundant in the winter and below detection levels during late spring and summer. Correlation analysis suggests that the production of the fronds of the giant kelp in this area was probably inhibited during the summer months by extremely low concentrations of nutrients. Internal nitrogen was exhausted approximately one month after a sharp decline in ambient nitrate concentration, and carbon reserves were formed. In the deeper bed of M. pyrifera, nitrogen was abundant all year round and the production of the fronds reflected the seasonal pattern of light or water temperature. The nitrogen content of the tissue probably did not drop below a level that limited production, and no internal carbon reserves were accumulated.     

Dynamics of shallow-water populations of Octopus dofleini

Octopus dofleini (Walker) is a common inhabitant of shallow subtidal communities in the Northeast Pacific. The abundances of octopuses at two sites monitored since 1977 have fluctuated greatly during this period. The populations reached high abundances in mid-summer and, during some years, mid-winter. The highest abundances at the two sites did not coincide. There was a constant influx of new octopuses into both study sites, with the greatest immigration occurring in early summer. The octopuses captured spanned a wide range of weights every month, with no clear size classes or seasonal trends in size. The weights of newly-captured octopuses, however, did differ between the sexes and seasons: males weighed more on the average than females, and male weight decreased from winter to fall while female weights did not change O. dofleini appeared to recruit throughout the year; the smallest octopuses occurred between May and November and the greatest number of small octopuses was found in July and August in most years. Females predominated at both study sites throughout the year. However, males predominated among octopuses caught in traps at nearby locations, suggesting that the skewed sex ratios were due to behavioral differences between the sexes.     

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.     

Differences between native and invasive<Emphasis Type="Italic"> Caulerpa taxifolia</Emphasis>: a link between asexual fragmentation and abundance in invasive populations

The tropical and subtropical marine green alga Caulerpa taxifolia has invaded several temperate regions throughout the world, including southeastern Australia. In this study, I examined how invasive C. taxifolia from temperate southeastern Australia differed from native C. taxifolia from subtropical Moreton Bay, Australia, in the traits proposed as being important to its invasion success: thallus size and density, levels of asexual reproduction (fragmentation) and total biomass. Against the prediction of a large size for invasive C. taxifolia, native populations from Moreton Bay had larger stolons and fronds than invasive populations. However, invasive populations consistently had much higher densities of stolons, fronds and fragmented fronds; and a greater biomass compared to native populations. Average densities at invasive sites exceeded 4,700 stolons and 9,000 fronds/m² and were as high as 27,000 stolons and 95,000 fronds/m², which are the highest reported for C. taxifolia anywhere. Average densities of fragmented fronds at invasive sites were as high as 6,000/m² and up to 45% of all stolons at invasive sites could be directly linked to asexual recruitment via fragmented fronds. Importantly, at invasive locations there was a strong association between asexual reproduction and abundance demonstrated by positive correlations between the density of fragmented fronds and total biomass. These findings are the first to describe quantitative differences between native and invasive C. taxifolia and to demonstrate a link between the high levels of asexual reproduction and high abundance in invasive populations. Although the causes and consequences of high levels of asexual reproduction remain to be explored, this study suggests that changes in demographic and life-history traits during the invasion by C. taxifolia into temperate habitats may contribute to its success there.Communicated by M. S. Johnson, Crawley     

The ecology of temperate salt-marsh fucoids. II. In situ growth of transplanted Ascophyllum nodosum ecads

Growth, in terms of length, weight, and number of branches and/or dichotomies, in transplanted specimens of Ascophyllum nodosum ecad scorpioides in a temperate salt marsh is described. The ecad scorpioides, when transplanted from its characteristic habitat on the mid-intertidal, Spartina alterniflora-dominated, marsh flats to a location near mean low-water developed characteristics normally associated with A. nodosum ecad mackaii. The growth of these plants was more rapid than those in the mid-intertidal region and was not affected by the shading of algal fronds by S. alterniflora. Unusually high temperatures and light intensities during the winter and spring months were major factors affecting growth in plants that were subjected to relatively long periods of tidal exposure. The presence of S. alterniflora during the summer months may act in a protective capacity for mid-intertidal ecad populations.This research was supported by research grants AG-375 and BO 38018 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).     

Photosynthesis in Codium fragile (Chlorophyta) from a Nova Scotia estuary: responses to desiccation and hyposalinity

Codium fragile ssp. tomentosoides from Caribou Harbour, an estuarine site in the southern Gulf of St. Lawrence, was extremely tolerant to stresses from desiccation and reduced salinity. Photosynthetic responses of both rhizomatous and erect growth forms were measured using pulse amplitude modulation (PAM) fluorometry of chlorophyll a fluorescence to determine effective quantum yield (Φ_PSII) and relative electron transport rate (rETR). After 5 h of desiccation, thalli lost 20% of their mass, but still showed high levels of Φ_PSII. Thalli survived for at least 6 h in freshwater, and showed virtually complete recovery of photosynthetic capacity within a few hours of return to full seawater. Immersion in 8 psu showed virtually complete recovery until the 24 h treatment period. Combining desiccation and salinity stresses produced a synergistic effect, but plants still showed strong recovery even after 86% dehydration and reimmersion in 16 psu. These results suggest that the photosynthetic physiology of Codium fragile is highly adapted to growth in estuarine conditions.     

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.     

Nitrogen limitation of Codium fragile ssp. tomentosoides as determined by tissue analysis

The critical nitrogen concentration of Codium fragile ssp. tomentosoides (van Goor) Silva was 1.90% on a dry weight basis. Internal nitrogen levels of thalli collected from three Rhode Island populations ranged from 0.75±0.08 to 3.72±0.08%. Internal nitrogen content was minimal in summer and maximal in winter, indicating nitrogen limitation during the summer and nitrogen storage during the winter. Part of this nitrogen storage appeared to support new growth of C. fragile in the spring. Carbon:nitrogen (C:N) ratios were inversely related to the internal nitrogen concentration. Low internal nitrogen content indicated nitrogen limitation better than C:N ratios because the C:N ratio could be influenced by both carbon and nitrogen metabolism.     

Biological traits of three closely related species of <Emphasis Type="Italic">Pycnoclavella</Emphasis> (Ascidiacea) in the Western Mediterranean

The reproductive cycles and abundance of the sympatric colonial ascidians Pycnoclavella brava, Pycnoclavella aurilucens and Pycnoclavella communis from two Northwestern Mediterranean sites over a period of 2.5 years are reported. The species showed some differences in their biological patterns. Gonad maturation and larval brooding took place during autumn and early winter for P. communis, during spring for P. aurilucens and during late winter and spring for P. brava. Summer was found to be the unfavorable season for all species, and an aestivation period occurred in P. communis and P. brava. Maximum abundance for the three species was observed during winter. Growth rates of the most abundant species, P. communis, were also monitored and found to peak during the initial phases of reactivation after aestivation. Growth rates decreased afterwards, becoming negative as water temperature started to increase after the winter minima. Reproductive activity and growth rates for P. communis displayed a temporal lag that suggested partitioning of resources to either reproduction or growth in this species.     

Seasonal stresses shift optimal intertidal algal habitats

We studied how the growth, reproduction, and survival of a common intertidal rockweed (Fucus distichus) varied across its tidal elevation at 14 sites around San Juan Island, Washington, USA in spring–summer and fall-winter seasons. We also measured a suite of environmental factors including temperature, light, emersion time, slope, fetch, and herbivory. To interpret the response of Fucus we included measurements of phlorotannins and carbon storage compounds (mannitol, laminarin). Growth and reproduction exhibited parallel patterns across tidal zones and sites. Tidal zone was a significant source of variation for many Fucus response variables, whereas variation between sites was high but not generally a significant factor explaining Fucus growth and physiology. Unexpectedly, the tidal zone in which Fucus achieved its highest growth and reproduction switched between seasons. High zone thalli grew and reproduced better than Mid zone thalli in fall but not in spring. This result can be explained by different combinations of factors influencing Fucus in each season. In spring, longer emersion times due to daytime low tides resulted in lower growth rates higher on the shore, likely due to carbon limitation. In fall during nighttime low tides, emersion and carbon limitation stresses were minimal. Overall, fall growth was lower than spring growth, but low fall light was not responsible. Instead, warmer average fall temperatures in the High zone apparently favored growth and reproduction relative to the Mid zone. In contrast, Mid zone thalli were subjected to more intense herbivory and hydrodynamic stress associated with wave exposure and steep substrata during the fall. At least for some seaweeds, living in the presumably more stressful high zone can actually confer higher integrated performance.     

Desiccation enhanced nitrogen uptake rates in intertidal seaweeds

Desiccation increased nitrate and ammonium uptake rates upon resubmergence in late summer populations of the intertidal macroalgae Gigartina papillata (C.Ag.) J.Ag., Enteromorpha intestinalis (L.) Grev., Fucus distichus L., and Pelvetiopsis limitata (Setch) Gardn. The ratio of nitrogen uptake rates in desiccated plants to rates in hydrated plants (controls) was correlated with the position of the species in the intertidal zone. Gracilaria pacifica Abbott., the species occurring at the lowest shore level, showed no enhancement of nitrogen uptake following desiccation. The high intertidal species such as P. limitata and F. distichus showed a two-fold enhancement of nitrate and ammonium uptake following more extensive desiccation (>30%) and continued uptake even following severe desiccation (50 to 60%). After the plants had been desiccated, the increase in nitrate uptake rates upon submergence lasted much longer than a similar enhancement of ammonium uptake. The duration of the enhanced nitrate uptake was similar to the time required for total rehydration but the uptake rates were not related to the state of rehydration. The potential contribution that this enhanced nitrogen uptake following desiccation could make to total nitrogen procurement for growth is discussed. The experiments were carried out in 1979 or 1980 and repeated in 1981.This paper is dedicated to Dr. R. F. Scagel on the occasion of his retirement for his outstanding contribution to phycology     

Seasonal changes in the chemical composition of Sargassum muticum

The alga Sargassum muticum (Yendo) Fensholt, a member of the Phaeophyceae, exhibits a growth cycle at Bembridge, Isle of Wight (UK), in which large annual fronds develop from much smaller perennating structures. Seasonal changes between April 1977 and June 1978 in the chemical composition of the alga were recorded. The content of inorganic ions, alginate and the dry weight: fresh weight ratio were relatively constant. Mannitol, laminaran and polyphenolic compounds were at their maxima in early summer (June/July), when the plant was undergoing rapid extension growth. In contrast, the protein content was highest in winter and early spring. It was concluded that the elongation phase which occurred in the spring was not supported by the use of stored polysaccharide reserves, since the amounts present were insufficient for this purpose and increased during the period of most rapid growth.