Growth of juvenile Macrocystis pyrifera (Laminariales) in relation to environmental factors

Instantaneous relative growth rates, (d^-1), were measured for juveniles of the giant kelp Macrocystis pyrifera transplanted to study sites in Southern California kelp forests between 1978 and 1982. Growth rates ranged from negative values (indicating loss of tissue) to 0.03 (doubling of total frond length every 19 d). Multiple regression analysis of growth versus irradiation, temperature, nitrogen concentration and amount of fouling revealed that all these factors had significant effects, together accounting for about 50% of the total variance. Elevated irradiation and nitrogen levels had strongly stimulatory effects (tissue nitrogen may have been more critical than ambient nitrogen for growth), while high temperature and fouling had strongly inhibitory effects. Irradiation was the most important factor influencing growth in 6 of the 8 transplant experiments. During these 6 experiments, the compensating irradiation level (below which there was no growth) was between 0.4 and 0.7 E m^-2 d^-1, and saturating irradiation was between 2 and 3 E m^-2 d^-1. During two of the experiments, growth was apparently limited by extremely high temperatures or low nitrogen levels. Quantum irradiation levels in the kelp forest were generally between the compensation and saturation levels. However, irradiation levels occasionally dropped below the compensation point for several months. Irradiation was occasionally low enough to limit the distribution of juvenile kelp by inhibiting growth, especially in the deeper portions of the kelp forest and under dense canopies formed by adult plants.     

Response of gametophytes of Ecklonia radiata (Laminariales) to temperature in saturating light

Gametophytes of Ecklonia radiata (C.Ag.)J.Ag. from two New Zealand locations with different field temperature ranges were exposed to temperatures of 5° to 26°C in saturating light. Plants from Goat Island Bay (Lat. 36° 16S, Long. 174°48E) grew in 9.3° to 25°C and reproduced in 9.3° to 24°C. There was no growth at 8°C and plants died at 26°C. Plants from the cooler location, Houghton Bay (Lat. 41°20S, Long. 174°40E), grew from 8° to 24°C and reproduced up to 15°C but not at 21.5°C. The plants did not grow at 6°C and died at 26°C. The timing of the first cell division and subsequent growth rate were retarded close to the upper and lower tolerance limits. Reproduction was a broad optimum of roughly 12° to 20°C. Within this range, fertile female gametophytes grown at lower temperatures had fewer, larger cells and thus fewer potential ova than those grown at higher temperatures.     

Nitrogen uptake kinetics in three year-classes of Laminaria groenlandica (Laminariales: Phaeophyta)

Nitrate and ammonium uptake rates were measured for three year-classes of the perennial macrophyte Laminaria groenlandica Rosenvinge, collected from nitrogen-depleted waters in Barkley Sound, British Columbia, Canada, in summer 1981. A time course of uptake rate revealed that ammonium uptake was high during the first hour and then decreased for all three year-classes; the opposite pattern was exhibited for the time course of nitrate uptake rate. Nitrate uptake rate increased linearly with nitrate concentration up to the highest level tested (60 M). The nitrate uptake rate of first-year plants was three times higher than second- and third-year plants; ammonium uptake rates showed similar patterns to those for nitrate. The interaction between nitrate and ammonium was examined for first-year plants. Nitrate and ammonium were taken up simultaneously and uptake rates were identical and equal to uptake rates when only nitrate or ammonium was present in the medium. Therefore, first-year plants are able to take up twice as much inorganic nitrogen per unit time when both nitrate and ammonium are present. First-year plants showed significant diel periodicity in ammonium uptake rates, whereas second- and third-year plants showed no periodicity in nitrate or ammonium uptake rates.     

Light and temperature requirements for growth and reproduction in gametophytes of Ecklonia maxima (Alariaceae: Laminariales)

The morphological and reproductive development of gametophytes of Ecklonia maxima (Osbeck) Papenf. has been studied in vitro under a wide range of light and temperature conditions, and the results related to the natural environment over the range of the distribution of the species. Initial vegetative growth was light saturated at 20 E m^-2 s^-1, and maximal at 17.5° and 20°C. Most rapid egg production corresponded to low cell number of female gametophytes, and this process was light saturated at 60 E m^-2 s^-1, and optimised at 15° and 17.5°C. Cell number was also low at lower temperatures, with reduced fertility rates. Sub-saturating irradiances and supraoptimal temperatures caused females to become filamentous, producing many more cells, and reducing reproductive rates. The final egg production per female was, however, greater in these sub-optimal conditions, and this phenomenon is interpreted as an ecological adaptation which improves survival prospects in conditions prevailing at the fringes of the depth and geographical distribution of the species. The optimal, and maximum (22.5°C) temperatures for reproduction are far higher than those of northern hemisphere Laminarians. E. maxima is very much a warm temperate organism, and this could have implications for marine phytogeographical studies on the west coast of southern Africa.     

Ammonium excretion in a temperate-reef community by a planktivorous fish,Chromis punctipinnis (Pomacentridae), and potential uptake by young giant kelp,Macrocystis pyrifera (Laminariales)

The blacksmith Chromis punctipinnis, an abundant planktivorous damselfish off southern California, USA, shelters along rocky reefs at night. While sheltered, blacksmiths excrete ammonium that could, in turn, be utilized by nearby benthic macrophytes. Laboratory experiments during the summer and fall of 1983 and 1984 indicate that ammonium excretion at night ranged from 18.1 mol h^-1 by a 8.5 g (dry) fish, to 89.1 mol h^-1 by a 27.3 g fish; excretion rates generally declined throughout the night. Field measurements at night indicate that ammonium concentrations were significantly higher in rocky crevices occupied by blacksmiths than in unoccupied shelters, and the ammonium level in one shelter dropped after a blacksmith was experimentally removed. Young kelp plants (Macrocystis pyrifera) are capable of taking up ammonium at night. Ammonium levels in chambers containing both a blacksmith and a young kelp plant were significantly lower than in chambers containing only a fish, and ammonium levels dropped in ammoniumspiked chambers that contained kelp plants. Nighttime ammonium uptake rates by young kelp plants, which averaged 1.6 mol g^-1 (dry) h^-1, were only slightly lower than those during the day. Daytime excretion by blacksmiths occasionally results in elevated ammonium levels in the water column. On two of six days, ammonium concentrations in midwater foraging aggregations were slightly but significantly higher than in upcurrent controls; since blacksmiths typically aggregate at the incurrent margin of kelp beds, the ammonium is swept downcurrent and may be utilized by large M. pyrifera that extend through the water column. Thus, the activities of blacksmiths may results in the importation of extrinsic, inorganic nitrogen to primary producers on temperate reefs.     

Allocation of organic material and energy to the holdfast,stipe, and fronds inPostelsia palmaeformis (Phaeophyta: Laminariales) on the California coast

Postelsia palmaeformis were collected from the lower intertidal at Pigeon Point, California, USA, in May 1987, and the proximate composition and allocation of energy to the various body components were determined. The holdfast and stipe have a proximate composition (% dry weight) of ca. 40% ash, 5.3% protein, 1% lipid, 2% soluble carbohydrate, and 55% insoluble carbohydrate. The fronds have a proximate composition of ca. 25% ash, 6.5% protein, 2% lipid, 3% soluble carbohydrate, and 65% insoluble carbohydrate. The energetic level was ca. 12 kJ g^-1 dry wt and ca. 19 kJ g^-1 ash-free dry wt. The relative proportion of three plant components varied, comprising 26, 39, and 35% wet wt and 20, 42, and 38% kJ for the holdfast, stipe, and fronds, respectively. A plant with a basal stipe diameter of 33 mm contains 114 g wet wt and 266 kJ. The maximal density found in May 1987 was 826 plants, 49 301 g wet wt, and 106 157 kJ m^-2.P. palmaeformis differs in these characteristics from another intertidal pheophyte,Durvillaea antarctica, that is found in a high-energy intertidal zone.     

Effects of ultraviolet radiation on different life cycle stages of the south Pacific kelps, Lessonia nigrescens and Lessonia trabeculata (Laminariales, Phaeophyceae)

The effects of exposure to ultraviolet radiation (UVR), 280–400 nm, in different life histories and development stages of the kelps, Lessonia nigrescens and L. trabeculata, collected in the south-east Pacific coast (30°S) were evaluated in the laboratory. Germination and viability (motile zoospores, settled spores), diameter of the primary cell of the gametophytes, percentage of female gametophytes, fertility and sporophytes production were measured after exposure to three radiation treatments (PAR; PAR + UVA; PAR + UVA + UVB). The effects of UVR in young sporophytes (diploid stage) were evaluated as changes in maximal quantum yield of chlorophyll fluorescence of photosystem II (PSII) (F _v/F _m). A significant decrease in all variables was observed for the treatment that included UVB (PAR + UVA + UVB) after 2 and 4 h of exposure, in relation to the control. The motile spores were more sensitive to UVR exposure compared to settled spores and gametophytes, suggesting that along with an increase in ontogenetic development; there is an increase in the tolerance to UVR. In addition, it was observed that early stages of the intertidal L. nigrescens were more tolerant to UVR compared to the subtidal L. trabeculata. These results allow initially to infer that UVR may be regarded as an important environmental factor influencing the upper limit of distribution of these species, mainly through its detrimental effects on the early stages of the life cycle.     

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.     

Seasonal variation in total and soluble tissue nitrogen of Pleurophycus gardneri (Phaeophyceae: Laminariales) in relation to environmental nitrate

Seasonal variations in tissue nitrogen (ethanol soluble nitrate and ninhydrin positive substances, as well as total nitrogen) of different thallus parts of Pleurophycus gardneri Setchell and Saunders were monitored simultaneously with ambient seawater nitrate from 1982 until 1984 in Bamfield, Vancouver Island, British Columbia, Canada. A trend of low, nearly zero levels in ambient nitrate typical for the area in late spring and early summer normally contrasts with average nitrate concentrations of 10 mol NO₃ ^- l^-1 in late fall and winter. Total nitrogen content was greater in the perennial thallus parts, stipe and holdfast than in the annual blade and peaked in fall and early winter. The longitudinal thallus distribution of nitrate revealed a distinct and significant concentration of nitrate in the haptera reaching at maximum 8% nitrate-N of the internal total nitrogen. Internal nitrate concentration ranged from 20 to 5 000 times the ambient nitrate concentration in the midrib, and from 40 to 3 100 times in the wing, while the range was greatest with 400 to 14 000 times in the haptera. P. gardneri contained at most about 7 mol NO₃ ^- g fresh wt^-1 in the blade, which corresponds to about 6% of total tissue nitrogen. Ninhydrin positive substances comprised the major portion of the soluble N pool in P. gardneri and showed a pronounced seasonality. Concentrations of ninhydrin positive substances ranged from 20 to 800 g N g fresh wt^-1 in the midrib and in the wing. In the stipe, ninhydrin positive substances varied from 180 to 2 200 g N g fresh wt^-1, and from 250 to 1 200 g N g fresh wt^-1 in the haptera. Evidence is given that (1) the perennial parts, stipe and haptera of P. gardneri contain the majority of nitrogen products independent of season and ontogenetic stage; (2) ninhydrin positive substances are the most abundant internal nitrogen constituents; (3) the low N values in the blade in summer suggest a nitrogen limited growth; and (4) nitrate may not be the predominant external nitrogen source.     

Morpho-functional patterns of photosynthesis and UV sensitivity in the kelp <Emphasis Type="Italic">Lessonia nigrescens</Emphasis> (Laminariales,Phaeophyta)

The photosynthetic responses of the south Pacific kelp Lessonia nigrescens of the coast of Valdivia, Chile (40°S), were investigated by exposing its different thallus parts, fronds, stipes and holdfasts, to UV radiation in the laboratory. Biologically effective doses (BED_{photoinhibition300}) between 400 and 800 kJ m⁻² were required for a 40% inhibition in photosynthesis under UVA+UVB radiation. At BED_{photoinhibition300} close to 250 kJ m⁻² (in treatments without UVB), the inhibition of photosynthesis did not exceed 20%. These UV doses were in the range of current daily doses measured in Valdivia on cloudless summer days. In general, exposure to UVB for periods longer than 12 h reduced photosynthesis, measured as maximal quantum yield (F _v/F _m) and electron transport. The fronds were the most UV-sensitive section of this alga, coinciding with the highest pigments contents and carbon fixation. Evidence of a photodamage was also seen. After a 48 h exposure to PAR+UVA+UVB, a decrease of F _v/F _m in the fronds was close to 41%, while in the stipes and holdfasts it was 12 and 18%, respectively. Although the thalli from the different size classes showed marked differences in their morphology and morphometry, no obvious differences in the UV tolerance of the fronds were detected. The results indicated that the UV-related responses are integrated in the suite of morpho-functional adaptations of the alga. Although the fronds are spatially more exposed to solar radiation than basal structures (stipes and holdfast), due their high turnover rate they may compensate better detrimental effects of UV. In contrast, stipes and the holdfast are key support structures characterized by low replacement rates and designed to confer hydrodynamic resistance to drag forces.     

Effects of the 1983-El Niño on growth and carbon and nitrogen metabolism ofPleurophycus gardneri (Phaeopyceae: Laminariales) in the northeastern Pacific

Growth and compounds of carbon and nitrogen of the northeast Pacific kelpPleurophycus gardneri (Laminariales, Laminariaceae) were assessed before, during and after the 1983-El Niño Souther Oscillation event. This 2-yr study provides evidence that reduced ambient levels of nitrate and phosphate from March until August/September, 1983, in combination with elevated seawater temperature, caused a significant impairment of growth. Blade surface area, blade length, total fresh weight, and dry weight on an areal basis were significantly reduced in 1983 compared to 1982 (P<0.01). They were also significantly smaller in 1984 than in 1983 (P<0.05). The C:N values obtained both in May and July 1983 were considerably higher than in 1982, while those in May 1984 were strikingly smaller than in both 1982 and 1983. Mannitol content of the mibrid in May 1983, exhibited the highest levels recorded during the 2-yr study period, with ca. 19% dry weight and significantly exceeded mannitol content measured in May 1982 (average: 13%) and in May 1984 (average: 8%). The concentration factor for internal vs external nitrate in the midrib was 700 x ambient in May 1982 and 1984 compared with 20 x in May 1983. Ninhydrin positive substances (NPS) averaged 117 mol g dry wt^-1 in May 1984 compared with ca. 50 mol in May 1982 and 1983. The pigment ratios chlorophyllc: chlorophylla and fucoxanthin: chlorophylla in May 1983 were similar to those obtained in 1982, but were significantly different in May 1984. It is suggested that the interactive effects of temperature stress and nutrient limitation in 1983 caused a disruptive influence on C- and N-metabolism ofPleurophycus gardneri, which still affected growth performance in 1984.     

Variability over space and time in the genetic structure of the winged kelp Alaria marginata

The genetic structure of Alaria marginata Postels & Ruprecht was investigated spatially and seasonally using amplified-fragment length polymorphisms (AFLP). Using one primer, 206 scoreable bands were produced. Individual plants that were separated by as little as a few decimeters to >100 km could be distinguished, and followed an isolation-by-distance model. Genetic similarity (average April data) ranged from 76% for patches (a few decimeters in diameter), to 71% for individual kelp stands (15 m across) and 67% for a group of stands separated by 185 km. Greater genetic similarity of patches occurred at a wave-sheltered site than at a wave-exposed site. The lower wave action may limit meiospore dispersal, resulting in patches of related sporophytes. Genetic similarities between A. marginata stands indicated a pattern of gene flow that is consistent with local currents. In one stand, genetic similarities were markedly different between seasons. This seasonal pattern may be the result of different age groups dominating the sampled stands over time. The genetic structure of A. marginata comprises local scale (patch and within-stand) heterogeneity and larger scale (between-stands) homogeneity. Received: 30 September 1998 / Accepted: 26 October 1999     

Response of Ecklonia radiata (Laminariales) to light at 15°C with reference to the field light budget at Goat Island Bay,New Zealand

Gametophytes of Ecklonia radiata (C.Ag.)J.Ag. grew in culture at 15°C under daily quantum doses ranging from 0.86 to 360 cE m^-2. Growth rates increased with quantum doses up to 40 cE m^-2 d^-1, then reproduction began and the relative growth constant declined while ovum release came earlier with increasing light up to about 100 cE m^-2 d^-1. Above 100 cE m^-2 d^-1 there were no consistent trends with increasing light, except that at the higher quantum doses, fertile female plants had fewer and larger cells and therefore fewer potential ova. Reproduction varied with daily quantum dose rather than with daylength. Given the same daily dose, plants grew fastest in low photon flux density, long daylength conditions. Gametophytes grown in the field developed at similar rates to those in culture. Gametophytes survived seven months of darkness at 10°C but died after one week of darkness at 20° to 23°C. Sunlight of 1 000 E m^-2 s^-1 was fatal to gametophytes and to sporophytes under 2 mm long after 10 to 15 min. Light budgets were prepared for plants growing at 7-and 15-m depths from 1976 to May 1980 in Goat Island Bay, New Zealand (Lat.36° 16S, Long. 174° 48E). Underwater light was measured under various environmental conditions. Relationships between transmission of light through the sea, data from diving visibility records and continuous surface meteorological records were studied. Approximations were made of the average percentage of surface light transmitted to 7 and 15 m over half-monthly periods. By applying these average transmittance values to the records of surface incident light, the average daily quantum doses were calculated. Light on open bottom in Goat Island Bay may sometimes be limiting for gametophyte reproduction in winter at 15-m depth. At depths less than 7 m, summer photon flux densities may reach damaging levels.     

A cell system for symplastic transport of photoassimilate between meristoderm and sieve elements in Alaria teniufolia

Prior to long-distance assimilate transport in the blades of Laminariales, photoassimilate from the meristoderm (source tissue) must first be transferred across the cortex before it can be loaded into the sieve elements. This radial assimilate transfer seems to be a symplastic parenchyma transport. The wings of Alaria tenuifolia Setchell are only a few cell layers thick and reveal a continuous cell system interconnecting the meristoderm with the sieve elements. This cell system probably provides an uninterrupted route for the symplastic transverse assimilate transfer and loading. The tissue topography of the wings, the anatomy of the composing cell types, and the micromorphology of the cellular network channelling assimilate to the sieve element system are described. The anatomical results are discussed with regard to the transport physiology of Laminariales in general and the possible mode of sieve element loading.     

祁连山青海云杉群落物种组成及α多样性垂直分布

通过对祁连山自然保护区青海云杉群落物种组成及α多样性垂直分布格局的研究,结果表明：祁连山自然保护区青海云杉群落内有维管植物25科51属96种,其中乔木4种,灌木29种,草本62种;植物种相对集中分布于海拔2 680～2 890 m的阴坡;随海拔梯度升高,群落内植物种数减少,且相邻样地的共有种数呈不明显的先减少而后增加的趋势,Margalef指数、Shannon-Wiener指数和Pielou指数呈降低趋势;不同林型的植物种的丰富度依次为：青海云杉混交林〉草类-青海云杉林〉苔藓-青海云杉林〉灌木-青海云杉林〉马先蒿-青海云杉林。     

Elemental composition (C,N,H) in larval and crab-1 stages of Pagurus bernhardus (Decapoda,Paguridae) and Carcinus maenas (Decapoda,Portunidae)

Analyses of individual content of carbon (C), nitrogen (N), and hydrogen (H) were carried out for all larval stages of Pagurus bernhardus and Carcinus maenas, and for newly metamorphosed crabs. Maximum range in total larval development is 12.8 to 165.8 g C, 3.2 to 35.1 g N, and 1.9 to 24.9 g H in P. bernhardus and 3.1 to 43.2 g C, 0.7 to 10.1 g N, and 0.4 to 6.3 g H in C. maenas. From these data energy equivalents were calculated. Maximum range in total larval life is 0.43 to 6.38 J ind. ^-1 in P. bernhardus and 0.1 to 1.49 J ind. ^-1 in C. maenas. There is a 32.4% mean loss of energy in P. bernhardus megalopa development; this seems to describe the normal developmental pattern in this stage. Biomass was determined as fresh and dry weight respectively. Individual dry weight is about 3.6 to 5.6 times higher in P. bernhardus (44 to 340 g) than in C. maenas (12 to 93 g) larvae.Contribution to research project Experimentelle marine Ökosystemanalyse sponsored by Bundesministerium für Forschung und Technologie, Bonn (Grant No. MFU-0328/1)     

Hydrolithon spp. (Rhodophyta, Corallinales) overgrow live corals (Cnidaria, Scleractinia) in Yemen

In Yemen, off the northwestern coast in the Gulf of Aden, the coralline algae Hydrolithon rupestre (Foslie) Penrose 1996 and H. murakoshii Iryu and Matsuda 1996 have been observed to overgrow and kill living Porites lutea Milne-Edwards and Haime, 1860. Similarly, Hydrolithon onkodes (Heydrich) Penrose and Woelkerling 1992 and H. rupestre were observed overgrowing Stylophora pistillata (Esper, 1797). Competitive interactions between P. lutea and H. murakoshii were monitored from 2006 to 2009 at two sites and showed an average linear growth of 8.3 (±1.9 SD) mm?year^?1 over the coral. The small polyps of S. pistillata and P. lutea combined with putative chemical compounds produced by Hydrolithon spp. are likely to allow the coralline overgrowth. Although corallines can locally kill coral tissues, the CCA/coral interactions do not seem to affect the overall live coral cover at the study sites.     

Comparative species richness,composition, and abundance of invertebrates in Caribbean seagrass (Thalassia testudinum) meadows (Panamá)

The results of a year-long study in which epibenthic invertebrates were collected monthly from seagrass (Thalassia testudinum) meadows along the Caribbean coast of Panamá and the Panamá Canal Zone are described in this paper. Differences in species composition and abundance among sites were primarily due to the proximity of surrounding habitats, especially coral reefs, which contain a number of species that utilize the seagrass meadows. In contrast to many previous characterizations of tropical marine habitats, important seasonal fluctuations in both species number and abundance took place at each of the sites. Data on breeding activity among several species of decapod crustaceans indicate year-round reproduction, although considerable seasonal differences occur in the percentage of ovigerous females. These interspecific differences in observed reproductive output may be explained by differences in life-cycle length, a factor not often considered in discussions of seasonal breeding patterns in tropical marine invertebrates. Overall species composition was qualitatively similar to that reported in comparable studies of tropical and subtropical seagrass meadows elsewhere, although caridean shrimp and xanthid orab species were reduced in number and total abundance were much lower than in previous studies.     

Growth and elemental composition (C,N, H) in Inachus dorsettensis (Decapoda: Majidae) larvae reared in the laboratory

Larvae of the spider crab Inachus dorsettensis were reared in the laboratory at constant 12 °C. Development lasted 8 to 10 d in the Zoea I, 10 to 12 d in the Zoea II and 14 to 20 d in the megalopa stage. During this time, larval growth was measured in samples taken every 2 to 4 d as dry wt (W), carbon (C), nitrogen (N), and hydrogen (H); energy content (E) was calculated from C. Biomass and energy (per individual) increased in each larval stage as a parabola-shaped function of age, which could be fitted by a power equation. C, H, and E show a higher percentage gain (relative to the initial values at hatching) than W or N, suggesting that proportionally more lipid than protein is accumulated during larval development. There are cyclical changes in the relative (per unit of W) biomass and energy figures, corresponding to the larval moult cycles: immediately after each ecdysis all these values decrease, presumably due to rapid uptake of water and minerals, then they increase again due to tissue growth and remain high until the next moulting occurs. Cyclical changes in the C/N ratio suggest that proportionally more lipid than protein is accumulated during the initial (postmoult) phase of the moult cycle, followed by a period of balanced or protein-dominated gain during the intermoult and premoult phases. These patterns of growth and elemental composition observed during the complete larval development and in single moult cycles of I. dorsettensis are compared with those described in the literature for other decapod species. This comparison suggests a high degree of similarity in biochemical composition and growth characteristics of larval decapod crustaceans.