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.     

Effect of seawater velocity on inorganic nitrogen uptake by morphologically distinct forms of Macrocystis integrifolia from wave-sheltered and exposed sites

In conditions of low water motion (<0.06 ms^–1), the availability of essential nutrients to macroalgae, and thus their potential productivity, may be limited by thick diffusion boundary-layers at the thallus surface. The ability of macroalgae to take up nutrients in slow moving water may be related to how their blade morphology affects diffusion boundarylayer thickness. For the giant kelp, Macrocystis integrifolia Bory, morphological measurements indicate that blades of plants from a site exposed to wave action are thick, narrow and have a heavily corrugated surface. In contrast, blades from a site with a low degree of water motion are relatively thin, with few surface corrugations and large undulations along their edges. The aim of our work was to test the hypothesis that morphological features of M. integrifolia blades from a sheltered site allow enhanced inorganic nitrogen uptake at low seawater velocities compared to blades with a wave-exposed morphology. The rate of nitrate and ammonium uptake by morphologically distinct blades of M. integrifolia, from sites that were sheltered from and exposed to wave action, were measured in the laboratory at a range of seawater velocities (0.01 to 0.16 ms^–1), between March and May 1993. For both sheltered and exposed blade morphologies, nitrate and ammonium uptake rates increased with increasing seawater velocity, reaching a maximum rate at 0.04 to 0.06 ms^–1. Uptake parameters V _max (maximum uptake rate) and U _0.37 (the velocity at which the uptake rate is 37% of the maximum rate) were estimated using an exponential decay formula. These parameters were similar for both blade morphologies, at all seawater velocities tested. Additional measurements suggest that the nitrogen status of M. integrifolia blades from wavesheltered and exposed sites were similar throughout the experimental period, and thus nitrogen status did not affect the rate of nitrogen uptake in these experiments. on the basis of these results, we conclude that blade morphology does not enhance nitrogen uptake by M. integrifolia in conditions of low water motion. Potential effects of diffusion boundary-layers on kelp productivity are discussed.     

Spatial and mass balanced trophic models of La Rinconada Marine Reserve (SE Pacific coast), a protected benthic ecosystem: Management strategy assessment

Steady-state, dynamic, and spatial models were constructed for the benthic system of La Rinconada Marine Reserve off northern Chile (SE Pacific coast). We examined data on biomass, P/B ratios, catches, food spectrum, consumption, and the dynamics of commercial and non-commercial populations using three theoretical frameworks: Ecopath, Ecosim, and Ecospace. The biomass of the scallop Argopecten purpuratus and the clam Tagelus dombeii were the most relevant compartments of the studied ecosystem. Among the carnivores, the functional crab group Cancer spp. was the most relevant. The Rhodophyta was the dominant macroalga compartment of the system. The results obtained using mixed trophic impacts (MTI) showed that the predatory snail Thais chocolata propagated higher magnitudes of direct and indirect effects on the other species or functional groups. The sea star Luidia magallanica and Rhodophyta had the least effects on the remaining compartments. According to the Ecosim estimates (increasing mortality by fishing), the scallop A. purpuratus had the highest impact on the other compartments. The Ecospace model showed similar qualitative and quantitative effects for changes in biomass under three different exploitation scenarios (by subsystems and globally). Nevertheless, the greatest changes were provoked by using the top-down control and the vulnerabilities estimated by Ecosim. System recovery times were highest with increased mortality of the asteroid L. magallanica and the carnivorous snail T. chocolata, suggesting that the sea star could be considered to be a top predator with a top-down control. The F_MSY estimated for the scallop A. purpuratus was close to the F_i originally entered in Ecopath, limiting the design and execution of an exploitation plan within ecologically sustainable boundaries. The situation was different (F_MSY ? F_i) for the other commercial species, making possible multi-species exploitation programs. The Ecospace trophic-spatially explicit model shows a similar pattern of direct and indirect effects generated when exerting exploitation separately by subsystems. Therefore, habitat rotation of fisheries is not justified.     

Ecological role of stomatopods (mantis shrimps) and potential impacts of trawling in a marine ecosystem of the southeast coast of India

Ecosystems are balanced by nature and each component in the system has a role in the sustenance of other components. A change in one component would invariably have an effect on others. Stomatopods (mantis shrimps) are common and ecologically important predatory crustaceans in tropical marine waters. The ecological role of mantis shrimps and potential impacts of trawling in a marine ecosystem were estimated using Ecopath with Ecosim (EwE) Version 5.0 software, by constructing a mass balanced Ecopath model of Parangipettai (Porto Novo) ecosystem. Based on fisheries information from the region, 17 ecological groups were defined including stomatopods. Both primary and secondary data on biomass, P/B, Q/B and diet composition were used as basic inputs. The mass balanced model gave a total system throughput of 14,756 t km⁻² year⁻¹. The gross efficiency of 0.000942 indicated higher contribution of lower food chain groups in the fishery though the mean trophic level was 3.08. The immature and developing stage of the ecosystem was indicated by the ratio of total primary production and total respiration (1.832) and the net system production (2643.30 t km⁻² year⁻¹). Key indices (flow to detritus, net efficiency and omnivory index), split mortality rates and mixed trophic impact of different ecological groups were obtained from the model. A flow diagram was constructed to illustrate the trophic interactions, which explained the biomass flows in the ecosystem with reference to stomatopods. Two temporal simulations were made, with 10 year durations in the mass balanced Ecopath model by using ecosim routine incorporated in EwE software. The effect of decrease in biomass of stomatopods in the ecosystem was well defined, in the first run with increase in stomatopod fishing mortality, and the group showed a high positive impact on benthopelagic fish biomass increase (129%). The simulation with increase in trawling efforts resulted in the biomass decline of different ecological groups as elasmobranchs to 1%, stomatopods to 2%, crabs and lobsters to 36%, cephalopods to 63%, mackerel to 78%, and shrimps to 89%. Present study warns stomatopod discards and further increase in trawling efforts in the region and it explained the need for ecosystem based fisheries management practices for the sustainability of marine fisheries.     

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.     

Representing mediating effects and species reintroductions in Ecopath with Ecosim

Ecosystem models play an important role in supporting ecosystem approaches to management. To improve the representation of how ecosystems work, ecosystem models should be able to represent mediating effects (e.g., habitat provision) that species provide to each other as well as species (re)introductions, both common situations that can strongly influence ecosystem dynamics. We examine how such processes can be incorporated into Ecopath with Ecosim (EwE), a widely used tool for represent aquatic ecosystems with the potential to support ecosystem-based management. We used the reintroduction of sea otters (Enhydralutris) to the west coast of Vancouver Island, British Columbia, Canada as a case study. The model demonstrates how to account for benefits provided by kelp forests by contributing to primary production, increased feeding areas and food availability through prey retention. It also demonstrates how the reintroduction and range expansion of sea otters can be represented in Ecospace, and the implications of these options.     

Assessing the carrying capacity of tilapia in an intertidal mangrove-based polyculture system of Pearl River Delta, China

A trophic model of an intertidal mangrove-based polyculture system in Pearl River Delta, China, was constructed using the Ecopath with Ecosim software. This polyculture system was chosen since it is the first integrated multi-trophic aquaculture (IMTA) system that was constructed on the basis of mangrove planting in China. The energy flows, ecosystem property, and carrying capacity of tilapia in the polyculture system were analyzed and evaluated. The results show the trophic level of 1.00 for primary producers and detritus to 2.85 for grass carp. The geometric mean of the trophic transfer efficiencies was 7.0%, with 7.2% from detritus and 6.8% from primary producers within the system. The ecosystem property indices show that this polyculture system has a high value of total primary production/total respiration (TPP/TR) and total primary production/total biomass (TPP/TB), together with low Finn's cycled index (FCI), Finn's mean path length (FML), and connectance index (CI), indicating that this system is at a development stage according to Odum's theory. The principal fish cultured in the system is tilapia, and mixed trophic impacts (MTI) show that tilapia has a marked impact on most compartments in this system, and the carrying capacity was found to be a tilapia culture biomass of 5.8 t ha⁻¹ in the system.     

Oviposition,larval abundance,in situ larval growth and recruitment of the herbivorous gastropodLacuna vincta in kelp canopies in Barkley Sound,Vancouver Island (British Columbia)

We investigated recruitment of the herbivorous gastropodLacuna vincta (Montagu, 1803) in the canopies ofMacrocystis integrifolia andNereocystis luetkeana beds in Barkley Sound, Vancouver Island (British Colombia), from 1987 to 1989. Four factors influencing intensity and patterns of recruitment were studied: (1) seasonality of oviposition, (2) larval abundance, (3) growth of larvae in the field and (4) larval settlement. Egg masses were abundant on low intertidal algae but were scarce in kelp canopies. Although egg masses could be found almost year-round, a distinct and intense period of oviposition occurred during winter and spring. Intracapsular development lasted 2.5 to 3.5 wk before planktotrophic veligers emerged. The duration of the planktonic period, 7 to 9 wk, was determined through an in situ study of cohorts ofLacuna spp. larvae present in the plankton between January and June 1988. The general timing of the onset of the spring peak recruitment period was predicted from these cohorts. Primary periods of recruitment ofL. vincta in the canopy occurred in April–May (average density up to 383.9 juveniles m^–2 blades), with a second period of lower intensity in the late summer—fall period. We observed similar trends between abundance of advanced larvae (> 500µm) in the plankton and recruitment rates in kelp canopies. Although adults were occasionally observed in the canopy, newly metamorphosed juveniles consistently dominated the habitat. The persistance of small juveniles (0.7 to 1.5 mm), rapid declines in density shortly after recruitment, and SCUBA observations of drifting individuals suggest that juveniles migrate to the under-canopy or low intertidal area after a brief period of growth on kelp blades.     

Species-specific defense strategies of vegetative versus reproductive blades of the Pacific kelps <Emphasis Type="Italic">Lessonia nigrescens</Emphasis> and <Emphasis Type="Italic">Macrocystis integrifolia</Emphasis>

Chemical defense is assumed to be costly and therefore algae should allocate defense investments in a way to reduce costs and optimize their overall fitness. Thus, lifetime expectation of particular tissues and their contribution to the fitness of the alga may affect defense allocation. Two brown algae common to the SE Pacific coasts, Lessonia nigrescens Bory and Macrocystis integrifolia Bory, feature important ontogenetic differences in the development of reproductive structures; in L. nigrescens blade tissues pass from a vegetative stage to a reproductive stage, while in M. integrifolia reproductive and vegetative functions are spatially separated on different blades. We hypothesized that vegetative blades of L. nigrescens with important future functions are more (or equally) defended than reproductive blades, whereas in M. integrifolia defense should be mainly allocated to reproductive blades (sporophylls), which are considered to make a higher contribution to fitness. Herein, within-plant variation in susceptibility of reproductive and vegetative tissues to herbivory and in allocation of phlorotannins (phenolics) and N-compounds was compared. The results show that phlorotannin and N-concentrations were higher in reproductive blade tissues for both investigated algae. However, preferences by amphipod grazers (Parhyalella penai) for either tissue type differed between the two algal species. Fresh reproductive tissue of L. nigrescens was more consumed than vegetative tissue, while the reverse was found in M. integrifolia, thus confirming the original hypothesis. This suggests that future fitness function might indeed be a useful predictor of anti-herbivore defense in large, perennial kelps. Results from feeding assays with artificial pellets that were made with air-dried material and extract-treated Ulva powder indicated that defenses in live algae are probably not based on chemicals that can be extracted or remain intact after air-drying and grinding up algal tissues. Instead, anti-herbivore defense against amphipod mesograzers seems to depend on structural traits of living algae.     

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.     

Assessing biomass gains from marsh restoration in Delaware Bay using Ecopath with Ecosim

The Delaware Bay ecosystem has been the focus of extensive habitat restoration efforts to offset finfish losses due to mortality associated with power plant water intake. As a result, a 45 km² or a 3% increase in total marsh area was achieved by 1996-1997 through the restoration efforts of the Public Service Enterprise Group (PSEG). To quantify the impact of restoration efforts on system productivity, an Ecopath with Ecosim model was constructed that represented all major components of the ecosystem. The model consisted of 47 functional groups including: 27 fish species, 5 invertebrate groups, 4 multi-species benthic groups, 6 multi-species fish groups, 3 plankton groups, 1 shorebird group and 1 marine mammal group. Biomass, abundance, catch, and demographic data were obtained from the literature or from individual stock assessments conducted for principal ecosystem components. A base Ecosim model was fitted to time series of key species in the Bay representing the period 1966-2003. To access the gains from marsh restoration, model simulations reflecting no restoration were conducted to estimate the productivity that would have been lost if restoration efforts had not occurred. The results indicated that restoration increased total ecosystem biomass by 47.7 t km⁻² year⁻¹. Simulations indicated increased biomasses across a wide range of species including important forage and commercially important species. The marsh restoration also significantly impacted ecosystem structure increasing the ratio of production-to-respiration, increasing system path length and decreasing the ratio of production-to-biomass.     

Integrative trophic network assessments of a lentic ecosystem by key ecological approaches of water chemistry, trophic guilds, and ecosystem health assessments along with an ECOPATH model

The objective of this study was to describe the trophic structure and energy flow in a lentic ecosystem in South Korea. Physicochemical water conditions were evaluated along with the reservoir ecosystem health using a multimetric IBI model. Nutrient analyses of the reservoir showed a nutrient rich and hypereutrophic system. Guild analysis revealed that tolerant and omnivorous species dominated the ecosystem. Tolerant fish, as a proportion of the number of individuals, were associated (R² > 0.90, p < 0.01) with TN and TP, the key indicators of trophic state in lentic ecosystems. The mean Reservoir Ecosystem Health Assessment (REHA) score was 19.3 during the study, which was judged as in ‘fair to poor’ condition. A trophic analysis of the reservoir estimated by the ECOPATH model shows that most activity in terms of energy flow occurred in the lower part of the trophic web, where there was intensive use of primary producers as a food source. Consequently, of the 10 consumer groups, nine fell within trophic levels <2.8. Trophic levels (TL) estimated from the weighted average of prey trophic levels varied from 1.0 for phytoplankton, macrophytes, and detritus to 3.25 for the top predator, Pseudobagrus fulvidraco. Our integrated approach to trophic network analysis may provide a key tool for determining the effects of nutrient influx on energy flow pathways in lentic ecosystems.     

Differential meiospore size and tolerance of ultraviolet light stress within and among kelp species along a depth gradient

 Kelp are differentially stratified along a gradient of UV exposure (as a function of water depth). The role of ultraviolet light in seaweed zonation has not been fully explored. This study found a significant meiospore size difference within and among the kelp species examined: Pterygophora californica Ruprecht (high to mid-subtidal), Macrocystis integrifolia Bory (high to mid-subtidal), Laminaria groenlandica Rosenvinge (high-subtidal), Alaria marginata Postels and Ruprecht (low-intertidal) and Hedophyllum sessile Setchell (mid-intertidal). This size difference was correlated with the depth distribution of adult plants, with the largest meiospores originating from shallow-dwelling adult kelp exposed to high UV light. Under UV stress in the laboratory, meiospores from adults growing in high-UV environments displayed greater germination and survival rates than the progeny of adult kelp occupying lower-UV environments. This suggested that in Barkley Sound, British Columbia, Canada, differential tolerance to UV (possibly determined by meiospore size) may limit the upper settlement position of kelp species and individuals. Tolerance to UV may be an important determinant of kelp zonation on rocky shores. Received: 16 June 1998 / Accepted: 6 January 2000     

Trophic relationships in the recently impounded Bagré reservoir in Burkina Faso

The trophic dynamics of Bagré reservoir which has been recently impounded in Burkina Faso was based on the data collected during 1997–1998 period using the Ecopath model and software. Total fish biomass is 22.63 t km⁻² and mainly represents trophic levels (TLs) 2 and 3. The trophic food chain is relatively long and the overall transfer efficiency is quite low. Grazing foodweb based on primary producers is prominent in the reservoir ecosystem and detritus plays a less significant role. Seasonal and long-term variations in water quality have significant influences on the lower TLs clearly showing a bottom-up functioning of the ecosystem. Environmental degradations, such as siltation occurring in the lake, suggest possible risks in limiting ecosystem productivity.     

Database-driven models of the world's Large Marine Ecosystems

We present a new methodology for database-driven ecosystem model generation and apply the methodology to the world's 66 currently defined Large Marine Ecosystems. The method relies on a large number of spatial and temporal databases, including FishBase, SeaLifeBase, as well as several other databases developed notably as part of the Sea Around Us project. The models are formulated using the freely available Ecopath with Ecosim (EwE) modeling approach and software. We tune the models by fitting to available time series data, but recognize that the models represent only a first-generation of database-driven ecosystem models. We use the models to obtain a first estimate of fish biomass in the world's LMEs. The biggest hurdles at present to further model development and validation are insufficient time series trend information, and data on spatial fishing effort.     

Floating kelps in Patagonian Fjords: an important vehicle for rafting invertebrates and its relevance for biogeography

Floating macroalgae are common dispersal vehicles for associated benthic invertebrates. In order to investigate the importance of kelp rafts for species dispersal in the Patagonian Fjord Region (PFR), the abundance and distribution pattern of floating kelps (Macrocystis pyrifera, Durvillaea antarctica) and of the invertebrate fauna associated with M. pyrifera were evaluated during austral spring of 2002–2005, 2008 and 2010. In the southernmost Magellan Region (MR), benthic M. pyrifera were additionally sampled to compare the community structures in both conditions. Floating kelps were abundant throughout the entire PFR, harbouring a diverse and abundant invertebrate fauna. The density of floating kelps increased towards the south. In the MR, a loss of species was observed between benthic and floating condition (e.g. decapods, echinoderms, several peracarid species), but a high diversity of organisms from all major phyla were observed on rafts. Throughout the PFR, the predominant rafting species belonged to the peracarids, molluscs and annelids, but the community composition differed between floating samples from the northern and southern zones of the PFR. Relative abundances of peracarids were higher in northern zones, whereas molluscs and annelids dominated in the southern areas. Species of the peracarid genera Peramphithoe, Gondogeneia, Bircenna and Limnoria were shared between all areas. The results suggest that kelp rafts not only contribute to local population connectivity in the PFR, but could also be an important dispersal vehicle for rafting species along the PFR in north–south direction, crossing the biogeographic boundary around Taitao Peninsula. Furthermore, the MR appears to be an important stepping stone for species dispersal via kelp rafting in the subantarctic region.     

Modeling changes in the coastal ecosystem of the Pearl River Estuary from 1981 to 1998

The coastal ecosystem of the Pearl River Estuary (PRE) has been overfished and received a high level of combined pollution since the 1980s. Ecopath with Ecosim was used to construct two ecosystem models (for 1981 and 1998) to characterize the food web structure and functioning of the ecosystem. Pedigree work and simple sensitivity analysis were carried out to evaluate the quality of data and the uncertainty of the models. The two models seem reliable with regards to input data of good quality. Comparing the variations of outputs of these two models aimed to facilitate assessment of changes of the ecosystem during the past two decades.The trophic structure of the ecosystem has changed with an increase in the biomass proportion of lower trophic level (TL) organisms and a decrease in top predator biomass proportion. All the indices of ecosystem maturity examined show that the system was in a more mature condition in 1981 than in 1998, although the system has been in a condition of stress due to anthropogenic disturbances, such as environmental pollution and habitat destruction since 1981. The ecosystem was aggregated into six and seven integral TLs in 1981 and 1998, respectively, using the trophic aggregation routine of Ecopath. Most of the total system biomass and catch took place at TL II and III in both years. But the distribution of the total system biomass and catch at different TLs changed with decreasing proportions in higher TLs in 1998. The mean transfer efficiency was 9.1% and 10.2% in 1981 and 1998, respectively.Comparative network analysis allowed quantification of the importance of direct and indirect trophic interactions among functional groups. Moreover, a method derived from the mixed trophic impact (MTI) analysis allowed estimating importance of groups in terms of “keystoneness” and identifying the keystone species in the two models over the past two decades. The results indicate that there were no clear keystone species in 1998 but two keystone species at medium trophic levels were identified in 1981. Moreover, organisms located at low trophic levels such as phytoplankton, zooplankton and benthic invertebrates were identified to have relatively high keystoneness in the ecosystem.     

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.