Effects of sediments on the development of Macrocystis pyrifera gametophytes

Rocky ocean bottoms are covered from time to time with small amounts of fine sediments. This material may interfere with the development of germling stages of Macrocystis pyrifera. Tests were performed by introducing sediments before and after spores were dispersed in culture dishes. 10 mg cm^-2 of sediment, enough to occlude the surface, prevents spore attachment, greatly reducing the probability of survival. Smothering of established germlings was severe at 108 mg sediment cm^-2. Water motion further reduced spore success where sediments were present, probably because of abrasive scour. Mechanisms of natural and human interference with M. pyrifera reproduction are suggested.     

Regulation of algal community development in a Macrocystis pyrifera forest

The effects of small and large-scale roughness, overstory development, competition for space with sessile animals, and grazing on algal community development in a subtidal Macrocystis pyrifera forest were examined using specially prepared concrete blocks as substrata. Variation in small-scale roughness (crevices and grooves in the order of 0.1 to 3 mm width and depth) had no significant effects on community composition. However, M. pyrifera colonization, algal diversity, and sessile animal biomass were higher near the upper horizontal edges of blocks and concrete prisms. This “edge” effect may result from a combination of increased spore and larval settlement and enhanced growth of plants and animals associated with the turbulent eddies formed around these obstructions. Natural and experimentally produced variations in the algal overstory demonstrated that the presence of an overstory can reduce algal diversity and cover beneath. Caging experiments suggest that predatory fishes and sea-stars indirectly affect the algal community by removing sessile animals (primarily bryozoans) which compete with the algae for space. The exclusion of grazers resulted in increased growth of Gigartina spp. Selective grazing on this genus may account for its reduced abundance in the study area.     

Interactive effects of light and temperature on sporophyte production in the giant kelp Macrocystis pyrifera

Laboratory experiments were conducted in 1984 to test for an interaction between irradiance and temperature in controlling sporophyte production in the giant kelp Macrocystis pyrifera. Gametophytes from plants in the San Onofre kelp forest were cultured under a combination of seven irradiance (2.9 to 30 E m^-2 s^-1) and ten temperature (11° to 20°C) levels. The minimum daily quantum dose for fertility of female gametophytes during the 42 d culture-period was 0.25 E m^-2 d^-1. The saturation quantum dose was approximately 0.60 E m^-2 d^-1. Temperatures between 11° and 20°C had little effect on the fertilityirradiance relationship, except at irradiations near the threshold and saturation levels. Cultures at 0.60 E m^-2 d^-1 had significantly lower fertility after 6 wk at 20°C than at the lower temperatures and there was a trend toward lower fertility at temperatures greater than 15°C in cultures at 0.25 E m^-2 d^-1. These differences were also reflected in the development time for female gametophytes. The 11°C cultures reached 50% fertility slightly faster and the 20°C cultures slower than cultures at intermediate temperatures.     

Pigment content and photosynthetic rate of the fronds of Macrocystis pyrifera

The Macrocystis pyrifera (L.) C. Ag. frond is here described in terms of chlorophyll a, fucoxanthin, chlorophyll c and photosynthetic rate. Pigment concentrations increased back from the apical meristem reaching a maximum after 2 to 3 m. Pigment concentrations were then generally constant throughout most of the length of the frond, finally decreasing again in the oldest parts of the frond with the exception of the sporophylls. Pigment ratios remained relatively constant throughout. Maximum net photosynthetic rates on a given frond showed a decrease with tissue age on both an area basis (1040 down to 463 nmol O₂ cm^-2 h^-1) and on a chlorophyll a basis, which was shown as half-saturation constants (quantum irradiance) which dropped on an area basis from 85 mol m^-2 sec^-1 at 4.5 m above the holdfast to 26 mol m^-2 sec^-1 at 15.5 m. Young sporophytes transplanted from the sea floor to the surface (12 m) tended to decrease pigment content, while those transplanted to the bottom tended to increase all pigments, but especially fucoxanthin. Photosynthetic rates, however, changed little on a unit area basis. The results of these data are considered in the light of recent work on photosynthetic units, tissue age effects and general adaptations of the M. pyrifera frond to its light environment.     

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     

Modelling the growth and harvest yield of the giant kelp Macrocystis pyrifera

The giant kelp Macrocystis pyrifera is one of the largest and fastest growing seaweeds and is dominant over large areas of the west coast of North America. A model of its growth has been developed which describes plant biomass and production over the course of a year as a function of environmental parameters which affect the light flux. Such parameters include water clarity, spacing between plants, bottom depth, latitude, harvesting activity, and photosynthetic response (P _max and I _k ). Model results for a standard set of conditions (latitude 33°N, 3 m plant spacing, water absorbance of 0.115 m^-1 and 12 m depth) yield a peak daily gross production of almost 6 g C m^-2 d^-1, peak daily net production of almost 3 g C m^-2 d^-1, and a peak specific growth rate of about 0.022 d^-1. Annual gross production for this case is 1 567 g C m^-2 yr^-1; annual net production is 537 g C m^-2 yr^-1. These values are comparable to those from field measurements. Size and timing of biomass and production peaks are affected by changes in the parameters describing the light field, with peaks usually occurring later in the year for more adverse circumstances. Inhigher latitudes, the seasonal variation is so extreme that the plant could not last the year at 53° N in 12 m of water, although it is able to survive the year in shallower water. Harvesting has severe effects on biomass and production. Model results suggest that light limitation is a very important constraint on kelp growth that should not be overlooked. This implies that differences in parameters describing two environments must be considered when comparing results obtained at different locales.     

Net primary production, growth, and standing crop of Macrocystis pyrifera in Southern California

Marine macroalgae are believed to be among the most productive autotrophs in the world. However, relatively little information exists about spatial and temporal variation in net primary production (NPP) by these organisms. The data presented here are being collected to investigate patterns and causes of variation in NPP by the giant kelp, Macrocystis pyrifera, which is believed to be one of the fastest growing autotrophs on earth. The standing crop and loss rates of M. pyrifera have been measured monthly in permanent plots at three sites in the Santa Barbara Channel, USA. Collection of these data began in June 2002 and is ongoing. Seasonal estimates of NPP and growth rate are made by combining the field data with a model of kelp dynamics. The purpose of this Data Paper is to make available a time series of M. pyrifera NPP, growth, and standing crop that is appropriate for examining seasonal and interannual patterns across multiple sites. Data on plant density in each plot and censuses of fronds on tagged plants at each site are also made available here. NPP, mass-specific growth rate, and standing crop are presented in four different metrics (wet mass, dry mass, carbon mass, and nitrogen mass) to facilitate comparisons with previous studies of M. pyrifera and with NPP measured in other ecosystems. Analyses of these data reveal seasonal cycles in growth and standing crop as well as substantial differences in M. pyrifera NPP among sites and years.     

Morphology and growth of the giant kelp Macrocystis pyrifera in New Zealand and California

At three sites (Papanui Inlet, Aquarium Point and Seacliff) on the Otago coast of New Zealand and at one site (Point Loma) off southern California, morphometric and shortterm growth measurements were made of single fronds of plants of Macrocystis pyrifera (L.) C. Ag. near the time of the autumn equinox, in 1977. Near the apex, internodes and laminae were shorter in wave-exposed Seacliff and California populations than in those of Papanui Inlet and Aquarium Point in strong currents, although lamina thickness was similar. In the immature region near the apex, linear relationships were apparent from double logarithmic plots of stipe distance from apex vs node number, internode fresh weight vs length (different in the two areas), lamina width vs length, fresh weight vs lamina area and immature lamina, pneumatocyst and internode fresh weight vs node number. Morphologically, Seacliff plants resembled those in California more closely than those in the other New Zealand populations. The relative growth rate in length of stipe internodes decreased linearly with the logarithm of the distance from the apex, but was less closely related to node number. Extrapolation gave zero growth at 1.6 m at Aquarium Point and 2.9 m in California, where the internode lengths coincided with a change in their weight/length relationship. Slow relative growth proximal to these points indicated a different type (mainly elongation). It is suggested that the first, apical, type of growth, being apparently unrelated to frond length, would be a useful measure of activity over short time-periods. The relative growth rate in lamina area (and probably volume), while also related to distance from the apex, showed a very different pattern, increasing slightly to a peak at 100 to 200 mm and declining to nil by 1 m at Aquarium Point and 2 m in California. Maximal lamina size was maintained for only 10 to 15 nodes at Aquarium Point. Calculated node initiation frequencies were 2.7 d at Papanui Inlet, 2 d at Aquarium Point, and 1 d in California. At the latter sites it took about 2 mo for laminae and 2^1/2 mo for stipe internodes to mature. Deduced relative growth rates in stipe tissue and pneumatocyst fresh weight decreased from 0.24 (log_e units per day) at the apex to 0.11 at 200 mm, where basal lamina tissue had a rate of 0.29, a doubling in just over 2 d. The production of tissue per frond was estimated as 25 and 36 g fresh wt d^-1 at Aquarium Point and in California, respectively.     

Nutrient-limited growth of juvenile kelp,Macrocystis pyrifera,during the 1982–1984 “El Niño” in southern California

The relative growth rates of juvenile Macrocystis pyrifera in southern California kelp forests were substantially reduced during the El Niño of 1982–1984. The lower growth rates were correlated with increased temperature and decreased nitrogen availability. Fertilization of juvenile plants with slow-release nitrogen-phosphorus fertilizer increased their growth rates to levels previously observed when temperatures were low and nutrient levels were high. The limitation in growth of M. pyrifera by levels of available nutrients during El Niño was in contrast to the usual limitation of growth by irradiance during non-El Niño years. Thus, there was a shift in the relative importance of factors controlling growth of juvenile M. pyrifera during El Niño.     

Morphological variations of Macrocystis pyrifera in the Falkland Islands in relation to environment and season

Some morphological characteristics of the giant kelp Macrocystis pyrifera in the Falkland Islands were studied from December 1985 until March 1987 in a shallow and relatively sheltered coastal zone and from December 1985 until January 1987 in a deeper offshore field exposed to swells. Seasonal fluctuations in lamina wet weight, density and form as well as pneumatocyst wet weight form and stipe density (i.e., wet weight per unit length) paralleled fluctuations in frond wet weight. Morphological differences between canopies of the giant kelp in the coastal zone and the offshore bed were probably mainly due to differences in water movement and depth between the two sites. Laminae and pneumatocysts of submersed-frond sites had different shapes than those of canopy-forming portions of fronds at the same sites, and their internodes were longer.     

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.     

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.     

Sediment load and timing of sedimentation affect spore establishment in Macrocystis pyrifera and Undaria pinnatifida

Although the frequency and magnitude of sedimentation often varies across coastal landscapes creating patches with different mean sediment loads, duration of sedimentation and rates of sediment resuspension, few studies have documented the emergent effects of spatio-temporal variability in sedimentation. Here, we conducted two laboratory experiments to evaluate such effects on the establishment of Macrocystis pyrifera and Undaria pinnatifida spores. In the first experiment, spore establishment was significantly affected by sediment load (the effective dose required for a 40 % reduction in establishment ranged between 16 and 60 mg sediment l^?1) and sediment regime (relative sedimentation occurring before spore settlement, ~3 times more sediment was required for 20 % reduction in spore establishment when sedimentation occurred after spore settlement). The second experiment demonstrated that the effects of sediment depended on sediment load (spore establishment was 2–4 times greater when sediment load was 200 mg l^?1 relative to 400 mg l^?1), variability in sedimentation (spore establishment was 1.36 times greater with variable than fixed sediment loads), repeated pulses of sedimentation (pulsed sedimentation decreased spore establishment by 59–91 % relative to a single sedimentation event) and timing of sedimentation relative to spore settlement (sedimentation before spore settlement decreased establishment by 51–95 % relative to sedimentation after spore settlement). These results have important implications for ecologists and resource managers attempting to predict the consequences of sedimentation, suggesting that it is not only important to consider sediment load, but also fine-scale temporal variability in sedimentation relative to key life-history events of the impacted organisms.     

Escape speeds of marine fish larvae during early development and starvation

Response rates to tactile stimulation and subsequent escape speeds were measured using a video-recording system during early development and starvation of fish larvae. The species studied included the yolk-sac larvae of Clyde and Baltic herring (Clupea harengus L.), cod (Gadus morhua L.), flounder (Platichthys flesus L.) and older larvae of Clyde herring. The proportion of larvae responding (response rate) was initially about 20 to 25% in herring and 35 to 40% in cod and flounder using a probe, but about 70 to 80% using the sucking action of a pipette in all species except flounder. Both response rates and escape speeds (mean and maximum) tended to peak 1 to 2 d before the PNR (point-of-no-return, when 50% of larvae are too weak to feed), then decreased slowly during further starvation. An inter-species comparison showed that the highest recorded mean escape speeds (measured over a period of 200 ms) and highest maximum escape speeds (over 20 ms) ranged from 5.7 to 8.6 BL/s (body lengths/s) and 12.1 to 16.1 BL/s, respectively. The larvae made directional responses away from the stimulus only when they developed and reached the feeding stage.     

Effects of an oil and gas-production effluent on the colonization potential of giant kelp (Macrocystis pyrifera) zoospores

Point sources of pollution (e.g. industrial and municipal outfalls) may produce ecological impacts at distant locations if pollutants affect dispersive propagules. We used laboratory experiments to determine how water-column exposure to produced water (=the aqueous fraction of oil and gas production that is typically discharged into coastal waters) influences the colonization potential of giant kelp (Macrocystis pyrifera) zoospores on the bottom. Zoospores were maintained in suspension at relatively low densities in 18-liter containers and exposed to one of five concentrations of produced water for varying amounts of time. Zoospore swimming generally decreased with increasing produced-water concentration and exposure duration; however, the specific pattern of decrease differed between experimental trials done on different dates. The effect of exposure duration on the ability of swimming zoospores to attach to plastic dishes placed on the bottom varied with produced-water concentration. Zoospores placed in produced-water concentrations of 1 and 10% showed a steady decline in their ability to attach with increased exposure; lower concentrations of produced water had no such effects. The percentage of zoospores that germinated after attachment varied tremendously with exposure duration and date of experimental trial. Zoospores that settled during the first 12 h after release had very poor rates of germination, indicative of a short precompetent period. Surprisingly, exposure of suspended zoospores to high concentrations of produced water during the first 12 h reduced this precompetent period and greatly improved germination success on the bottom. The magnitude of this enhancement, however, varied among dates. The results suggest that adverse effects of discharging produced water on planktonic zoospores of giant kelp would most likely be limited to the immediate vicinity of the outfall.     

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.     

Synchronized breeding events in sympatric marine invertebrates: role of behavior and fine temporal windows in maintaining reproductive isolation

While breeding synchrony among conspecifics is increasingly well understood with regards to the reproductive success of vertebrate and invertebrate taxa, the occurrence of simultaneous multispecies breeding events remains intriguing. The fairly recent discovery of mass annual spawnings in reef corals has provided a first glimpse at putative strategies of reproductive isolation during such events. However, the mechanisms and advantages of same-day heterospecific breeding are still poorly understood and have not yet been investigated in non-coral taxa with different life history strategies. In an effort to bridge this gap, we investigated spawning periodicity and synchrony among 26 sympatric species of free-spawning, capsule-laying, and brood-protecting macroinvertebrates belonging to six different phyla. Twenty-four of these species released gametes or larvae between early March and late April. We analyzed the events over fine temporal scales to test the hypothesis that breeding activities were not random in time or relative to each other. We found that the two main reproductive pulses followed a lunar periodicity and that consistent species- and gender-specific modulations in the timing of spawning occurred during same-day episodes involving up to six free-spawning species. Mass spawning accounts from the literature were reviewed and compared. This work suggests that many species participate in synchronous heterospecific spawning events either because they respond to the same environmental cues or rely on cross-cueing and that reproductive isolation is favored by species-specific circadian patterns, spawning behaviors and cross-gender signaling.     

Chemotactic effects of nutrients on spores of the kelps Macrocytis pyrifera and Pterygophora california

Fertile Macrocystis pyrifera (L.) C. Ag. and Pterygophora californica Rupr. were collected in California, USA in 1987 to 1988. Spores of the kelps exhibited both positive and negative chemotaxis to a variety of chemical nutrients. Chemotaxis was measured by counting the number of spores that swam into flattened capillary tubes with the chemical relative to the number that swam into control tubes. Video-motion-analysis also showed that P. californica spores swam towards a nitrogen source more often than they swam away. Similar chemotactic effects were observed in both 2 and 8 h-old preparations. M. pyrifera spores swam towards nitrate, ammonium (1 to 90 M), glycine, aspartate iron (1 m), boron, cobalt, and manganese. Negative chemotaxis was elicited by ammonium (1 000 M) and iron (45 M). Neither phosphate nor zinc had significant effects. P. californica spores were attracted by nitrate, ammonium, phosphate, and boron. Negative chemotactic effects were recorded with iron (45 M) and manganese. Iron (1 M), cobalt, and zinc had no effect. It is suggested that chemotactic behavior is an adaptation which allows the kelp spores to find and settle in microhabitats suitable for gamatophytic growth and reproduction.     

Interactive effects of pH,temperature and light during ammonia toxicity events in Elodea canadensis

Increased nutrient loading threatens many freshwater ecosystems. Elevated temperatures may increase the sensitivity to eutrophication in these ecosystems. Higher concentrations of possibly toxic reduced nitrogen (NH _x ) in the water layer may be expected as production and anaerobic breakdown rates will increase. Apart from temperature, NH _x and its effect on aquatic macrophytes will also depend on pH and light. We examined the interactive effects of NH _x , temperature, pH and light on Elodea canadensis in a full factorial laboratory experiment. Results demonstrate that high NH _x and high temperature together with low pH and low light causes the strongest toxic effects regarding relative growth rate and leaf tissue mortality. The adverse effects of high temperature and low light are most likely caused by increased metabolic activity and reduced photosynthesis, respectively. Severe toxicity at low pH compared to high pH can be ascribed to the ability of E. canadensis to induce a specialised bicarbonate-concentrating pathway at high pH, resulting in much higher carbon availability, needed for detoxification of NH _x . We conclude that NH _x toxicity will become more pronounced under higher temperatures, but that effects on aquatic macrophytes will strongly depend on pH of the water layer and specific metabolic adaptations of different species.     

Ocean acidification and temperature rise: effects on calcification during early development of the cuttlefish Sepia officinalis

This study investigated the effects of seawater pH (i.e., 8.10, 7.85 and 7.60) and temperature (16 and 19 °C) on (a) the abiotic conditions in the fluid surrounding the embryo (viz. the perivitelline fluid), (b) growth, development and (c) cuttlebone calcification of embryonic and juvenile stages of the cephalopod Sepia officinalis. Egg swelling increased in response to acidification or warming, leading to an increase in egg surface while the interactive effects suggested a limited plasticity of the swelling modulation. Embryos experienced elevated pCO₂ conditions in the perivitelline fluid (>3-fold higher pCO₂ than that of ambient seawater), rendering the medium under-saturated even under ambient conditions. The growth of both embryos and juveniles was unaffected by pH, whereas ⁴⁵Ca incorporation in cuttlebone increased significantly with decreasing pH at both temperatures. This phenomenon of hypercalcification is limited to only a number of animals but does not guarantee functional performance and calls for better mechanistic understanding of calcification processes.