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.     

Effects of grazing by two species of sea urchins (Strongylocentrotus franciscanus and Lytechinus anamesus) on recruitment and survival of two species of kelp (Macrocystis pyrifera and Pterygophora californica)

We studied the effects of grazing by two species of sea urchins on two species of kelp (Macrocystis pyrifera and Pterygophora californica) in the San Onofre kelp bed in southern California from 1978 through 1981. Both red sea urchins, Strongylocentrotus franciscanus, and white sea urchins, Lytechinus anamesus, were abundant and lived in aggregations. The purple sea urchin (S. purpuratus) was rare at the study site and was not studied. The aggregations of red urchins were either relatively small and stationary (for over 3 yr) or relatively large and motile (advancing at about 2 m mo^–1). Both stationary and moving aggregations were observed at the same time, and within 100 m of one another. Stationary aggregations of red urchins probably subsisted mainly on drift kelp and had no effect on kelp recruitment or on adult kelp abundance. In contrast, red sea urchins in large, motile aggregations or fronts ate almost all the macroalgae in their path. The condition of their gonalds indicated that red urchins in fronts were starved relative to red urchins in the small, stationary aggregations. Large, motile aggregations developed after 2 yr of declining kelp abundance (probably due largely to storms). We propose that a scarcity of drift algae for food results in a change in the behavior pattern of the red urchins and thus leads to the formation of large, motile aggregations. The aggregations of white urchins, which occurred along the offshore margin of the kelp bed, were large, but relatively stationary. The white urchins rarely ate adult kelps, but grazed extensively on early developmental stages of kelps and evidently prevented seaward expansion of the bed. The spatial distribution of both types of red urchin aggregations appeared to be unrelated to predation pressure from fishes or lobsters.Please address all requests for reprints to the senior author at his present address.     

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.     

Nuclear events during early development in gametophytes of Macrocystis pyrifera,and the temporal effects of a marine contaminant

The time course of DNA synthesis in developing haploid gametophytes of the giant kelp Macrocystis pyrifera was determined, and the effects of arsenic (As) on the temporally distinct nuclear events, DNA synthesis and subsequent nuclear division/translocation, were investigated to establish which of these specific events may be disrupted by this contaminant. Experiments were carried out on material collected from kelp beds near Santa Barbara, California from 1993–1994. Timing of DNA synthesis was determined during development by use of the fluorochrome, DAPI (4,6-diamidino-2-phenylindole), and single-cell microspectrofluorometry. Zoospores, which result from meiosis, had already undergone two rounds of DNA synthesis at the time of release. The developing gametophytes underwent an additional two rounds of replication of DNA by 16 h of development, and following the first nuclear division/translocation, the gametophyte contained eight times the minimum DNA level throughout subsequent development. Both DNA synthesis and nuclear division/translocation, were found to be inhibited by As. Phosphate enrichment reduced the inhibitory effects of As on division/translocation of the nucleus, supporting the hypothesis that As interferes with phosphorylation. Gametophytes were more severely affected by As under light conditions, as opposed to dark, suggesting that photosynthesis may be more sensitive than dark metabolism.     

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.     

Effect of bryozoan colonization on inorganic nitrogen acquisition by the kelps Agarum fimbriatum and Macrocystis integrifolia

The effect of bryozoan colonization on inorganic nitrogen acquisition by Agarum fimbriatum Harv. and Macrocystis integrifolia Bory., collected from the west coast of Vancouver Island, British Columbia, Canada, was examined in laboratory experiments during June and July 1992. Pieces of kelp blades that were completely covered on one side by the bryozoans Lichenopora novae-zelandiae Busk or Membranipora membranacea, L., or uncolonized (clean treatment), were used to estimate the rate at which nitrate and ammonium were removed from the surrounding seawater. In addition, the rate of ammonium excretion by bryozoans isolated from their associated kelp was measured and also estimated from the results of the uptake experiments. Values obtained were used to estimate the contribution of ammonium excreted by bryozoans to the total amount of inorganic nitrogen available to the associated kelp. Both bryozoan species reduced the ability of the associated kelp to remove nitrate and ammonium from seawater but provided a source of ammonium to the kelp through excretion. The nitrogen status of colonized and clean kelp disks was determined from the ratio of total particulate carbon to total particulate nitrogen (C:N ratio). The C:N ratios for A. fimbriatum colonized with either L. novae-zelandiae or M. membranacea were similar (C:N=12 to 14), and differences between colonized and clean treatments were not significant. For A. fimbriatum, therefore, the C:N ratio indicates that this species was not nitrogen limited at the time of the present study. In contrast, both colonized and clean disks of M. integrifolia were nitrogen limited, but colonized disks (C:N=19) were significantly less limited by nitrogen than clean disks (C:N=29). Results are discussed in relation to the different environments inhabited by both kelp species and are consistent with the hypothesis that ammonium excreted by bryozoans was an important source of inorganic nitrogen to M. integrifolia, but not to A. fimbriatum, at the time of the study.     

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.     

Geographic variation in polyphenolic levels of Northeastern Pacific kelps and rockweeds

Brown algal polyphenolic compounds are secondary metabolites whose functions may include protecting plants from pathogens or damage by UV radiation, and deterring feeding by herbivores. We present here the first analysis of spatial variation (at scales from tens of meters to hundreds of kilometers) in concentration of these compounds in two orders of brown algae from the northeastern Pacific Ocean. In kelps (order Laminariales), variation among sites was significant in only 25% of species examined and was consistent within families (high in the Alariaceae and low in the Laminariaceae and Lessoniaceae). In rockweeds (order Fucales, family Fucaceae), site variation was high in three of four species examined. Both the proportion of high polyphenolic kelp species and the magnitude of spatial variation within species from both kelps and rockweeds were much higher than would have been predicted from previous studies in other regions. In one kelp (Laminaria groenlandica), significant differences between sites occurred at scales of only tens of meters. No latitudinal clines were observed. Differences in phenolic concentrations of kelps spanned nearly an order of magnitude in one species, Hedophyllum sessile. Phenolic levels were significantly higher in members of the Fucales than the Laminariales, but showed no significant differences between intertidal and subtidal species. Received: 22 July 1996 / Accepted: 26 October 1998     

Nutrient stimulation of spore settlement in the kelpsPterygophora californica andMacrocystis pyrifera

FertilePterygophora californica Rupr. andMacrocystis pyrifera (L.) C.Ag. were collected in California, USA, from 1987 to 1989. Settlement activity of the spores was stimulated by nutrients, but this was not constant over time. Nutrients had no effect on the settlement ofP. californica spores between 2 and 18 h after sunrise, but settlement activity was stimulated by a nutrient mixture between 20 and 24 h after sunrise (14 to 18 h after release). Settlement activity inM. pyrifera spores was unaffected at 2 to 3 h after release, but settlement activity was significantly stimulated from 5 to 12 h after release. A variety of individual nutrients significantly stimulated settlement in 8 to 9 h oldM. pyrifera spores: ammonium, nitrate, glycine, phosphate, manganese-EDTA (18µM), borate, ferrous iron-EDTA, and ferric iron-EDTA. Spores also settled readily in unenriched artificial seawater, and nutrient-stimulated settlement rates were usually 150% of the unenriched control levels. Neither EDTA alone, cobalt-EDTA, nor manganese-EDTA (2µM) had significant effects onM. pyrifera spore settlement. The effects of time and of several individual nutrients on spore settlement activity are different from previously reported chemotactic effects of nutrients onP. californica andM. pyrifera spores. It is suggested that nutrient settlement-stimulation is mechanistically different from nutrient chemotaxis. However, like chemotaxis, settlement stimulation is probably an adaptation which increases the likelihood of spore settlement in microhabitats suitable for subsequent growth and reproduction of gametophytes.     

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     

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.     

Effect of boundary layer transport on the fixation of carbon by the giant kelp Macrocystis pyrifera

The uptake of inorganic carbon into the thallus of Macrocystis pyrifera (L.) C. Ag. requires first that the inorganic carbon pass through the water medium to the plant surface. This transfer of inorganic carbon to the thallus must take place through a boundary layer. Experiments in water tunnels indicate that the boundary layer adjacent to the M. pyrifera blade may be turbulent in water speeds as low as 1 cm sec^-1. Photosynthetic output of the blade can be increased by a factor of 300% by increasing water speeds over the blade surface from 0 to 4 cm sec^-1. This is consistent with a decrease in the thickness of the boundary layer. Above 4 cm sec^-1, the assimilation of carbon was limiting. The assimilation of carbon is generally known to follow Michaelis-Menten-like kinetics. Combining the two uptake steps into an overall model of carbon uptake agrees well with photosynthetic data obtained from M. pyrifera under varying conditions of water speed and bicarbonate concentrations in the laboratory. The ecological and morphological consequences of these findings are discussed.     

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

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

In vivo accumulation of radioactive polonium by the giant kelp Macrocystis pyrifera

The in vivo accumulation rate of ²¹⁰Po by Macrocystis pyrifera (L) Ag. has been determined by the tagging of specific parts of young, growing lamina in La Jolla kelp beds and their subsequent collection for ²¹⁰Po analysis. The rates of accumulation for observed and growth-corrected data were, respectively, 0.78x10^-9 pCi/cm² sec (1.73x10^-9 dpm/cm² sec) and 1.17x10^-9 pCi/cm² sec (2.60x10^-9 dpm/cm² sec). Invert, but biologically foulable, glass-slide surfaces exposed to the kelp bed environment yielded ²¹⁰Po accumulation rates of 0.64x10^-9 pCi/cm² sec (1.42x10^-9 dpm/cm² sec), slightly less than those of M. pyrifera. Distinction is made between observed net accumulation rate and gross rates or total flux. Several factors contributing to the final net accumulation rate are discussed.     

In situ water motion and nutrient uptake by the giant kelp Macrocystis pyrifera

Rates of NO ₃ ^- uptake by individual blades of Macrocystis pyrifera (L.) C. Agardh were measured at different flow rates in the laboratory. Dissolution rates of hemispherical, plaster buttons attached to the blade surface provided a relative measure of flow rates over blades used in uptake experiments and also over intact blades of adult kelp plants in situ (Laguna Beach, California, USA; 1981). Laboratory results indicated that uptake was saturated at a flow rate equivalent to 2.5 cm s^-1 current velocity. Flow rates over intact blades in situ always exceeded this uptake saturation level. Wave surge and movement of plant surfaces relative to the surrounding water provided sufficient flow to saturate uptake, even in a dense kelp canopy during low-current and calm sea-state conditions.     

Quantitative and qualitative observations on Laminaria dentigera and other subtidal kelps of southern Kodiak Island,Alaska

Observations were made and quantitative samples of non-canopy kelps were collected, using SCUBA, at 9 subtidal sites off southern Kodiak Island in May, 1976. At a 10th site, only observations were made. The most abundant species in the quadrats were Laminaria dentigera, L. yezoensis, Pleurophycus gardneri, Agarum cribrosum, and Alaria marginata. We found fertile plants of all these species. L. dentigera dominated at all sampled sites except one within a bed of Nereocystis luetkeana, where P. gardneri was dominant. L. yezoensis was dominant at the site which was visited but not sampled. The average wet weight of non-canopy kelps in the 55 samples was 12 kg m^-2, and the quadrat range was 2 to 35 kg m^-2. The reported ranges of P. gardneri and Alaria marginata are extended westward from Montague Island (147°22W) to Bumble Bay, Kodiak Island (154°43W), where A. marginata was found at a depth of 8.5 m.     

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.     

Dispersal of faunal and floral propagules associated with drifting Macrocystis pyrifera plants

The potential of drifting Macrocystis pyrifera kelp for transporting associated animals and plants long distances around the southern oceans was assessed by anchoring kelp holdfasts off the Tasmanian coast in 1985, monitoring the turnover of organisms, and relating species survival to water-transport times and species geographic distributions. Although most of the common animal species and approximately half of the plant species associated with Tasmanian M. pyrifera holdfasts were still present on kelp holdfasts after 191 d at sea, very few of these species have been recorded from New Zealand. It therefore seems unlikely that M. pyrifera plants with intact holdfasts are presently drifting to New Zealand. Drifting kelps probably become negatively buoyant in the Tasman Sea because dissolved nitrate concentrations are insufficient for normal plant growth. Moreover, even if some kelp plants do drift to New Zealand it is possible that their holdfasts rapidly disintegrate in the open ocean because of the abundance of the boring isopods Phycolimnoria spp. in Tasmanian holdfasts. In contrast to the restricted distributions of Tasmanian holdfast-inhabiting species, most of the identified species collected from M. pyrifera holdfasts at subantarctic Macquarie Island also occurred 5 000 km west at Kerguelen Island. Because of the extensive ranges of many subantarctic species, the good probability of survival of epifaunal species on drifting kelps, and the high surface-water nitrate concentrations and low holdfast-densities of Phycolimnoria spp. in the higher latitudes, it is likely that M. pyrifera-mediated transport of faunal and floral propagules has recently occurred, and is probably presently occurring, in subantarctic waters.     

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.