Is kelp detritus a good food for suspension feeders? Effects of kelp species, age and secondary metabolites

The food quality of detrital particles derived from three species of kelps was evaluated in a laboratory feeding experiment utilizing two species of suspension feeders, the serpulid polychaete Pseudochitonopomaoccidentalis and the mussel Mytilustrossulus. Fresh and aged kelp particles were also evaluated, and growth in all treatments was compared to growth on ad libidum phytoplankton rations. Fresh particles from Laminariagroenlandica, aged particles from Agarumfimbriatum and Alariamarginata, and mixed phytoplankton promoted the highest growth rates in both consumers. Growth was inversely related to total polyphenolic concentration in the fresh kelp particles. The increase in quality of both Agarumfimbriatum and Alariamarginata particles with age corresponded with a rapid loss of polyphenolic secondary metabolites and an increase in total nitrogen. Received: 26 November 1996 / Accepted: 9 January 1997     

Red algae as hosts for endophytic kelp gametophytes

We observed kelp gametophytes endophytic in the cell walls of 17 species of red algae from the San Juan Islands, Washington, USA. Host algae were collected primarily from three sites dominated by different kelp assemblages, including (1) a subtidal site dominated by Agarum fimbriatum Harvey, (2) a second subtidal site dominated by Nereocystis luetkeana (Mertens) Postels et Ruprecht, and (3) a floating dock on which the dominant kelp species were Alaria marginata Postels et Ruprecht, Laminaria groenlandica Rosenvinge, and Costaria costata (C. Agardh) Saunders. Most hosts were filamentous [e.g. Pleonosporium vancouverianum (J. Agardh) J. Agardh, Callithamnion acutum Kylin], or polysiphonous [e.g. Polysiphonia paniculata Montagne, Pterosiphonia dendroidea (Montagne) Falkenberg]; however, similar endophytes were also observed in membranous or bladed red algae [e.g. Membranoptera platyphylla (Setchell et Gardner) Kylin, Polyneura latissima (Harvey) Kylin]. Dozens to hundreds of separate kelp gametophytes were associated with single host thalli. In many cases, the gametophytes developed conspicuous oogonia raised above the thallus surface on characteristic stalks. Presumed zygotes developed through typical embryonic stages into juvenile sporophytes. We argue that the endophytic habit plays an important role in the biology of kelp gametophytes. Received: 5 May 1998 / Accepted: 6 May 1999     

Effect of solar ultra-violet radiation on the kelp Ecklonia radiata

During spring and summer, 1982–1986, experiments were carried out near Marmion Reef, Western Australia. In summer, nearly 30% of the surface solar ultraviolet radiation (280 to 400 nm) penetrates offshore waters to 5 m depth. Experimental removal of the mature Ecklonia radiata kelp canopy in summer results in tissue damage, photopigment destruction, reduced growth, and low survivorship of subcanopy kelp sporophytes. These effects do not occur with canopy removal in winter. Laboratory experiments revealed that the UV component of radiation, rather than intense photosynthetically active radiation, was responsible for the inhibition of growth and photodamage. UV radiation probably affects survival of the settlement stages of E. radiata sporophytes, thus excluding them from otherwise suitable substrata in shallow waters. UV radiation is implicated in the reduction of canopy productivity in summer.     

An annual carbon budget for the kelp Laminaria longicruris

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

Ecotypic differentiation in light-related traits of the kelp Laminaria saccharina

Light-related traits were compared for Laminaria saccharina Lamour. collected from three habitats in Maine and New York, USA, with different ambient light regimes. Light-level, expressed as a proportion of surface irradiance (I₀), ranged from 0.04 to 0.32 I₀ in the shallow habitat, but rarely exceeded 0.04 I₀ in the deep and turbid habitats. Juvenile sporophytes collected from each habitat in April, 1985, were grown at four acclimation light-levels (0.065, 0.12, 0.26, and 0.54 I₀) in a common-garden, laboratory experiment. Photosynthesis vs irradiance (PI) parameters, light-harvesting characteristics, and rates of carbon-assimilation and growth were determined for each group of plants. The results indicated that ecotypic differentiation had occurred among the three kelp populations. Photosynthetic capacity (P_max) and photosynthetic efficiency () were generally highest for plants from the turbid habitat, lowest for deep plants, and intermediate for shallow plants. These differences were largely attributable to variations in light-harvesting characteristics. The nature and magnitude of photoacclimation responses also differed among populations. Population differences in photosynthetic parameters resulted in different rates of C-assimilation and growth by plants from shallow, deep, and turbid habitats. Predictions of in situ growth rates indicated that the severity of light-limitation and, therefore, the adaptive significance of efficient light-utilization vary among the three populations. It is concluded that ecotypic differentiation in light-related traits is important to the broad environmetal range of L. saccharina.     

In situ exudation of phlorotannins by the sublittoral kelp Ecklonia radiata

Exudation of phlorotannins (polyphenolics) was measured in situ from the sublittoral kelp Ecklonia radiata at two locations near Sydney, New South Wales, Australia during 1992–1993. Minimally disruptive techniques were used in which individual plants were enclosed within clear plastic bags in order to concentrate exudates. Rates of exudation from E. radiata were low relative to most previous studies, with a mean rate (averaged across four seasons and two sites) of 5.5 g g (dry wt)^-1 h^-1. Exudation was greatest in summer and least in winter at one site, but there were no seasonal differences at a second, more protected site. There was no measurable diurnal variation in exudation rates. Exudation after a period of heavy storms was not significantly different from exudation during calm weather, but severe physical damage to the kelp did increase exudation. Our results suggest that exudation of phlorotannins in temperate Australian waters may be less ecologically important than has been suggested for coastal systems in the Northern Hemisphere.     

Effects of petroleum products on bull kelp (Nereocystis luetkeana)

Although research has been conducted on the effects of oil on the giant kelp Macrocystis pyrifera, no similar studies have been completed on bull kelp, Nereocystis leutkeana, the dominant kelp in Washington State, British Columbia, and Alaska. The effects of three petroleum products [diesel fuel, intermediate fuel oil (IFO), and crude oil] were tested before and after weathering on N. luetkeana. Whole plants were exposed to petroleum product for 4 or 24 h and then transferred to the field; observations on the condition of the plants were made daily for 7 d. In addition, controlled bioassays were performed to measure the effects of petroleum exposure on net photosynthetic rate (NP) and respiration rate (R), using light-and dark-bottle techniques. These experiments verified the susceptibility of N. luetkeana tissue to the damaging effects of direct exposure to several oil types. The 4 h exposures to weathered diesel and unweathered IFO, and 24 h exposures to unweathered and weathered diesel and IFO resulted in moderate to severe damage to kelp tissue (i.e., clearly delineated bleached line accompanied by tissue necrosis). Weathered diesel was more toxic than unweathered diesel. The most severe damage to bull kelp was concentrated at the meristematic zone (junction of stipe and bulb) where new tissue growth occurs. Petroleum type significantly affected stipe and blade NP, R, and NP:R ratios. Diesel treatments had a greater negative effect on NP than did the IFO treatments. Based on these experiments, the relative ranking of the damaging effects of petroleum treatment on bull kelp are weathered diesel>unweathered IFO>unweathered diesel>weathered IFO>unweathered crude>weathered crude.     

Distribution and abundance of marine nematodes on the kelp Macrocystic integrifolia

The species and distribution of nematodes on the kelp Macrocystic integrifolia Bory, in the Bamfield region of Barkley Sound, British Columbia, Canada were examined. Nine species (belonging to six families) of nematodes were found on the kelp blades. Three species (Monhystera disjuncta, M. refringens and Prochromadorella neapolitana) comprised 91–99% of the nematode fauna and occurred in all monthly samples from July 1978 to November 1979. The three dominant species exhibited seasonal density differences. P. neapolitana occurred mostly in the summer, M. refringens abundance peaked from July to October and M. disjuncta was relatively abundant throughout the year. The other species contributed little to the overall abundance and distribution patterns. All three species occurred in greatest abundance on the lower and middle blades of M. integrifolia in the deep end of the kelp bed. Very few individuals occurred on the upper blades. Nematode distribution on M. integrifolia appeared to be related to blade age and the associated food sources on the blade.     

Some biological interactions affecting intertidal populations of the kelp Egregia laevigata

Near Santa Barbara, California (USA), the large laminarian kelp Egregia laevigata (Setchell) occurred from the lower intertidal zone to subtidal depths. In the intertidal zone there was a large recruitment of E. laevigata in the spring. The kelp were largely excluded from a zone in the lowest intertidal zone where the surf grass Phyllospadix sp, grew, but were abundant on all other rock surfaces. Experiments demonstrated that, while neither grazers nor sessile organisms significantly reduced recruitment, E. laevigata of the previous year-class did so. There were also interactions among E. laevigata of the same year-class, expressed as a density-dependent mortality of very small algae, and as faster growth rates and greater number of branches of kelp at low densities. These processes tended to make both numbers and biomass of E. laevigata uniform. The mortality rates of large E. laevigata were so high that, in some locations, no kelp survived for more than 8 months. E. laevigata was, therefore, essentially an annual and opportunistic species in the intertidal zone.     

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     

Temperature ecotypes near the southern boundary of the kelp Laminaria saccharina

Effects of temperature on survival, growth, and photosynthesis were compared for two USA populations of Laminaria saccharina Lamour. One population was located in New York State, near the southern latitudinal boundary of the species in the western North Atlantic. This southern boundary population was exposed to ambient temperatures 20°C for about 6 wk each summer. The second population was located in Maine, toward the center of the latitudinal range of the species, and was rarely exposed to temperatures>17°C. sporophytes from the New York (NY) population exhibited greater tolerance of high temperature than plants from the Maine (ME) site. Juvenile sporophytes from the two sites had similar rates of survivorship and growth at temperatures below 20°C, but showed different responses at 20°C in laboratory experiments. NY plants survived and grew for 6 wk at 20°C. ME plants showed negative growth during wk 2 and 100% mortality during wk 3. NY and ME plants held in situ at the NY site during June to September, 1985, also exhibited differential survivorship when ambient temperatures exceeded 20°C. Results of photosynthesis and dark respiration measurements on NY and ME plants grown at various temperatures suggested that the high-temperature tolerance of NY plants was attributable to their ability to maintain positive daily net C-fixation at 20°C. The high-temperature tolerance of the NY plants appeared to be due to genetic adaptation and is probably crucial to the persistence of the species near its southern boundary.     

Changes in sea urchin populations after the destruction of kelp beds

In St. Margaret's Bay, Nova Scotia, Canada, there are large areas in which sea urchins (Strongylocentrotus droebachiensis) have eliminated beds of kelp (Laminaria spp.). Sites were identified where destruction of kelp beds had taken place 1, 2, 3, 3.5 and 4 years ago. With increase of time since kelp disappearance, the sea urchins showed decreased growth rate, reduced gonad size, but an increase in numbers resulting from high recruitment rates in the first two years after kelp bed destruction. These sea urchin populations, by their browsing, effectively prevent the regeneration of kelp. There is, as yet, no evidence of the sea urchin populations being starved out to allow the kelp to return.     

Modeling kelp forest distribution and biomass along temperate rocky coastlines

We address the global deficit of data describing kelp forest distribution, relative covers and biomass by testing the ability of species distribution models to predict these attributes at locations where data are currently limited. We integrated biological ground truth data with high-resolution environmental datasets to develop generalized additive models that accurately predict the structure of Laminaria forests within the Bay of Morlaix (48°42′42″N, 3°55′40″W). Forest distribution and proportional covers were predicted using water depth, light availability, wave exposure and sediment dynamics. The biomass of individual kelp species was modeled by supplementing these same variables with measures of seafloor slope and benthic position. Biomass predictions for Laminaria digitata and Laminaria hyperborea contrast the physiological tolerances of these species to light and wave exposure gradients. As a direct management output, we produced high-resolution maps (25 m² grids) that closely match independent field data and provide vital information for marine spatial planning.     

Photosynthetic characteristics of giant kelp (Macrocystis pyrifera) determined in situ

Rates of net photosynthesis and nocturnal respiration by individual blades of the giant kelp Macrocystis pyrifera (L.) C. Agardh in southern California, were determined in situ by measuring oxygen production in polyethylene bags during spring/summer of 1983. Mature blades from different depths in the water column exhibited different photosynthetic characteristics. Blades from the surface canopy (0 to 1 m depth) exhibited higher photosynthetic capacity under saturating irradiance and higher photosynthetic efficiency at low irradiances than blades from 3 to 5 or 7 to 9 m depths. Saturating irradiance was lower for canopy blades than for deeper blades. Canopy blades showed no short-term photoinhibition, but photosynthetic rates of deeper blades were significantly reduced during 1 to 2 h incubations at high irradiances. Results of 1 to 2 wk acclimation experiments indicated that differences between photosynthetic characteristics of blades from different depths were primarily attributable to acclimation light conditions. Vertical displacement of blades within the kelp canopy occurred on a time-scale of 1 min to 1 h. Blades continually moved between the unshaded surface layer and deeper, shaded layers. Vertical movement did not maximize photosynthesis by individual blades; only a small proportion of blades making up a dense surface canopy maintained light-saturated photosynthetic rates during midday incubations. The relatively high photosynthetic rates exhibited by canopy blades over the entire range of light conditions probably resulted from acclimation to intermittent high and low irradiances, a consequence of vertical displacement. Vertical displacement also reduced the afternoon depression in photosynthesis of individual canopy blades. The overall effect of vertical displacement was optimization of total net photosynthesis by the kelp canopy and, therefore, optimization of whole-plant production.     

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.     

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.     

Changing lobster abundance and the destruction of kelp beds by sea urchins

In a study area in Nova Scotia, Canada, abundance of the lobster Homarus americanus decreased by nearly 50% in 14 years. The lobster is a major predator of sea urchins, and during the past 6 years the sea urchin Strongylocentrotus droebachiensis has destroyed 70% of the beds of Laminaria spp. in the area. Implications for management are discussed.