Influences of abundance,behavior, species composition,and ontogenetic stage on active emergence of meiobenthic copepods in subtropical habitats

The emergence of meiobenthic copepods in subtidal sand and seagrass sites in Tampa Bay, Florida (USA), was investigated on nine dates from March 1983 to August 1984. Numbers of emerging copepods were dependent on habitat, date and diel sampling period. Greater numbers typically emerged in seagrass sites, and differences between habitats may be related to lower copepod abundances in sand sites. Both copepod abundance and behavior significantly affected numbers emerging among dates. An increase in emergent behavior alone resulted in consistently greater numbers emerging during postsunset periods. Over 30 species in 15 families were found to emerge. Total numbers of emerging copepods were affected by the sporadic presence and seasonal behavior of certain species. Numbers of the predominant speciesParadactylopodia brevicornis, Tisbe furcata, Parategastes sp.,Mesochra pygmea, Laophontid sp., andHarpacticus sp. emerging depended on differences in both species abundance and behavior. The emergent behavior ofParategastes sp.,Harpacticus sp., andT. furcata also was influenced by total copepod densities in the sediment. Juvenile copepods constituted only a small proportion of the total numbers emerging. The consistent postsunset entry of between 10³ to 10⁵ copepods m^?2 into the water column in subtidal sand and seagrass habitats will contribute to the increased probability of copepod dispersal and reassortment of the benthic community, while providing a pathway for benthic-pelagic exchange.     

Survival of copepods passing through a nuclear power station on northeastern Long Island Sound,USA

About 70% of the copepods entering the cooling water system of a nuclear power plant on northeastern Long Island Sound (USA) are not returned to the Sound in the effluent. Copepod mortalities are caused by the mechanical or hydraulic stresses of passage, although our experimental design could not determine whether heat or chlorination could cause mortality in the absence of mortality induced by hydraulic stress. After passing through the power plant, copepods sink rapidly (ca. 2.5 times faster than controls). This leads to an increase in concentrations of copepods suspended in the deep water (25 to 30 m) of the effluent pond. About half of the live copepods collected at the discharge and held in situ died within 3.5 days, and 70% died within 5 days, whereas only 10% of those from the intake died in 5 days. About 60% of the copepods observed suspended in deep water in the pond were dead. The copepod mortality caused by the power plant reflects the loss in secondary production occurring below about 270×10³ m² of sea surface in Long Island Sound annually. This loss represents a reduction of about 0.1% in the annual secondary production over a 333 km² area of Long Island Sound adjacent to the power plant. Highest losses occurred during the spring (April, 1.4×10⁶ g dry weight), the lowest in autumn (November, 45.8×10³ g). If the same copepod loss rate exists for all power plants in Long Island Sound, then secondary production in 1.69×10⁶ m², or 0.05% of the total copepod production may be lost annually. A comparison of the surface outflow from Long Island to Block Island Sound with the water entrained through Millstone Unit One, and the 70% copepod loss rate in the latter area, indicates that Unit One eliminates about 0.1 to 0.3% of the copepod production in eastern Long Island Sound. This calculation compares favorably with losses computed from production data.     

Resting eggs of zooplankton (Copepoda and Cladocera) from the Kiel Bay and adjacent waters (southwestern Baltic)

Resting eggs of four species of calanoid copepods and three species of cladocerans were collected from sediments up to 5cm depth from the Kiel Bay and adjacent waters in the southwestern Baltic Sea during April–May 1994. All but one species of cladoceran was successfully hatched/reared in the laboratory. In the Kiel Bay, egg abundances varied from 1.8x10⁵ to 7.4x10⁵ m^-2. Hatching success of copepod eggs collected from all depths was high (49 to 94%), but was 0 to 79% for cladoceran eggs. Darkness did not seem to affect hatching. Eggs found in the 4 to 5 cm layer of sediment were estimated to be about 15 yr old, showing the presence of an egg bank in the Baltic. Formation of resting eggs may be a genetic trait acquired during the ice ages.     

Feeding ecology of Spinocalanus antarcticus, a mesopelagic copepod with a looped gut

Spinocalanus antarcticus, an abundant mesopelagic copepod in polar seas, has a greatly elongated and looped midgut, contrary to most other copepod species. The total gut length is 1.77, 1.86 and 1.90 times the total body length in adult females, CV and CIV, respectively. Gross morphology of the midgut is similar in all copepodite stages and adults. It is described here from specimens collected in the Arctic Ocean. In stratified samples from the deep Amundsen and Makarov Basins S. antarcticus showed a clear preference for the depth layer between 100 and 500 m. Generally, the guts were packed with material, but most of it was impossible to identify. In most specimens the digestive tract was filled with undefined detritus particles ("detritus balls"). They were almost spherical, heterogeneous organic aggregates of 40-100 µm diameter, with small clay-sized mineral flakes imbedded. Mineral particles in the size range of 1-10 µm were found in large quantities in the guts of many specimens. Cysts of Chrysophycea and dinoflagellates and fragments of dinoflagellates, diatoms, tintinnids and radiolarians, as well as skeletons of silicoflagellates, were rather rare; some animal remnants were also found. A high carbon/nitrogen ratio (8.9) and very high lipid content (54% of dry weight) indicated a very good nutritional state. The adaptive significance and possible feeding strategy of this deep-water copepod is discussed.     

Mobility and adsorption of the triketone herbicide mesotrione in the soil of corn crops

The triketone herbicide mesotrione has been applied pre‐emergence at the dose of 150 g a.i. ha^?1 on corn fields grown within the same period of time at different sites on clay, loam, sandy loam or sandy soils. During the crops and one month after the corn harvest, the mobility of mesotrione has been measured in the 0–20 cm surface soil layer, soil samples being taken in the 0–2,2–4,4–6,6–8, 8–10, 10–15 and 15–20 cm surface soil layers. During the first month after the treatment, mesotrione remained in the 0–2 cm surface soil layer. Thereafter, mesotrione was at a higher concentration in a 2 cm‐thick soil layer which progressively moved down in the clay, loam and sandy loam soils. In the sand soil, mesotrione moved down as a uniform diffusion, its concentration being similar in all the 2 cm‐thick layers of the 0–10 cm soil surface layer. Low mesotrione residues (6–10 ug kg^?1 dry soil) attained the 10–15 cm layer faster in the clay and sand soils than in the loam and sandy loam soils. Mesotrione was not detected in the 15–20 cm soil layer of the loam and sandy loam soils, but well in the sand and clay soils. In the soils of the corn crops mesotrione thus mainly remained in the 0–10 cm surface soil layer. This low mobility and depth of penetration conjugated to the rate of mesotrione soil degradation explain why there was no movement of mesotrione toward the deeper soil layers. The adsorption of mesotrione was greatest on the soils recently treated with organic fertilizers, and having a loam or sandy loam texture. The adsorption coefficients explained the lower mobility and the greatest persistence of mesotrione in these soils.     

Photosensitivity of the calanoid copepod Acartia tonsa

The light intensity and spectral sensitivities of the calanoid copepod Acartia tonsa Dana were determined by measuring phototactic responses. Adult females displayed only positive phototaxis. The dark-adapted copepod, which possesses a single naupliar eye, perceived light at intensities as low as 2.8x10¹¹ photons m^-2 s^-1. The action spectrum for positive phototaxis had no clear maxima but rather showed a broad range of greatest sensitivity from 453 to 620 nm. This sensitivity encompassed those wavelengths that are maximally available at the depth where the copepod is found during the day. This spectral overlap, coupled with the finding that the copepod requires light cues for nocturnal vertical miration, suggests that broad spectral sensitivity is an adaptive mechanism to maximize light intensity sensitivity during migration.     

An individual-based model to study the reproduction of egg bearing copepods: Application to Eurytemora affinis (Copepoda Calanoida) from the Seine estuary,France

Limited empirical studies have elucidated the daily egg production and associated reproductive processes of egg bearing copepod. Herein, we present an individual-based model which constitutes a realistic representation of the reproduction in egg bearing copepods. The model has been parameterized using an extensive set of experimental data obtained from the literature and from the laboratory and field experiments on the estuarine copepod Eurytemora affinis. The proposed model takes into account the adult female longevity, the clutch size and interclutch duration, which is a function of egg maturation time and latency time required by the female after egg hatching to produce a new clutch. The embryonic development time and hatching success are also taken into account. The effect of temperature on the means and variances of above-mentioned reproductive parameters has been also incorporated. A multi agent system based generic platform “Mobidyc” has been used to generate and calibrate the model. The model demonstrates the reproductive parameters of females of E. affinis which is validated through individual based experiments. Temperature specific simulations provide a dynamical explanation of temperature effect on the cumulative egg production. The daily survival principally affects the number of clutches produced per female during its life span. The results obtained in the present study by combining temperature and survival effects reveal the relatively greater importance of the first factor on the daily egg production of egg-carrying copepods. The present model is generic and hence easily applicable to other animals with comparable reproductive strategy.     

Effects of prey concentration,prey size,predator life stage,predator starvation,and season on predation rates of the carnivorous copepod Euchaeta elongata

Adult females of the large carnivorous copepod Euchaeta elongata Esterly were collected from 1977 to 1980 in Port Susan, Washington, USA. Predation rates of the adult females increased with increasing prey abundance when fed the following 4 sizes of copepods: adult females of Calanus pacificus (average prosome length [PL] of 2 650 μm), adults of Aetideus divergens (PL of 1 560 μm), adult females of Pseudocalanus spp. (PL of 1 060 μm), and nauplii of C. pacificus (PL of 410 μm). Saturation feeding levels were reached when adult females of the predator were fed the small adult copepod, Pseudocalanus spp. Maximum biomass ingested of this small copepod was more than the maximum amount ingested of the larger copepods. Predation rates of the predatory copepodids at Stages IV and V also increased with increasing concentration of the 1 060 μm (PL) prey. High feeding rates exhibited by both adults and copepodids at Stage V of the predator indicate their importance as sources of mortality on populations of small copepods. Ingestion efficiency E _i (prey wholly consumed [prey attacked]^-1) varied as follows: adults of E. elongata were more efficient than copepodids of E. elongata; adults were more efficient than copepodids when ingesting smaller prey; starved adults were more efficient than fed ones; and both adults and copepodids were more efficient at low food concentrations. For adults of E. elongata, there were no marked seasonal variations in predation or respiratory rates that would represent acclimatory responses; however, small adults obtained during winter were more efficient at ingesting prey than were the larger adults gathered in summer. This seasonal variation in the efficiency of ingestion may be a useful indicator of physiological state: high E _i values could indicate that predators are starving in winter, and low E _i values could indicate that predators are satiated in summer.     

O3浓度升高对麦田土壤碳、氮含量和酶活性的影响

近地层O₃作为全球最重要的大气污染物之一,其对作物的生长发育、土壤酶活性、土壤碳、氮的影响机制已成为人们关注的重要问题。采用开顶式气室(OTCs)法模拟研究O₃浓度升高对冬小麦土壤碳、氮含量和酶活性的影响。结果表明,O₃浓度升高导致麦田0~10 cm和10~20 cm土层的全碳(TC)和全氮(TN)含量呈现出下降的趋势。O₃浓度升高对土壤酶活性也有影响。在冬小麦灌浆期,O₃胁迫可促进土壤脱氢酶活性提高。当O₃浓度为120 nL·L^-1时,0~10 cm、10~20 cm和20~40 cm土层的脱氢酶活性分别比对照处理提高59.4%、51.5%和22.2%。O₃胁迫对土壤转化酶活性的影响随着冬小麦生长期和土壤采样深度的不同而发生变化。在冬小麦拔节期,O₃处理对不同土层脲酶活性的影响没有达到显著差异水平,但是在灌浆期,20~40 cm土层的脲酶活性随着O₃浓度的增加而提高,在120 nL·L^-1浓度O₃处理下脲酶活性比对照处理提高24.6%。在O₃胁迫条件下土壤转化酶活性与土壤全碳含量、土壤脲酶活性与土壤全氮含量均呈现出显著的正相关关系。     

Biological weighting of ultraviolet (280–400 nm) induced mortality in marine zooplankton and fish. II. Calanus finmarchicus (Copepoda) eggs

The copepod Calanus finmarchicus Gunnerus is a key component of the planktonic food web in the Gulf of St. Lawrence, Canada. In this region, productivity-determining biophysical interactions occur in the upper 0 to 30 m of the water column. The eggs and nauplii of C. finmarchicus are found in this layer. Measurements of the diffuse attenuation coefficients for solar ultraviolet-B radiation (280 to 320 nm, UV-B) at various locations in this region indicated maximum 10% depths (the depth to which 10% of the surface energy penetrates) of 3 to 4 m at a wavelength of 310 nm. This represents a significant percentage of the summer mixed-layer water column: organisms residing in this layer are exposed to UV-B radiation. Laboratory experiments using a Xenon-arc-lamp based solar simulator revealed that C. finmarchicus embryos exposed to UV-B exhibited high wavelength-dependent mortality. The strongest effects occurred under exposures to wavelengths below 312 nm. A significant percentage of nauplii hatched from eggs exposed to these wavelengths exhibited malformations indicative of errors in pattern formation during embryogenesis. At the shorter wavelengths (<305 nm), UV-B-induced mortality was strongly dependent on cumulative exposure. The biological weighting function (BWF) derived for UV-B-induced mortality in C. finmarchicus eggs is similar to that reported for naked DNA. This suggests that the UV-B-induced mortality effect on C. finmarchicus embryos is a direct result of DNA damage. There was no evidence of a detrimental effect of ultraviolet-A radiation (320 to 400 nm). Calculations based upon the BWF indicate that, under current noon surface irradiance, 50% of C. finmarchicus eggs located at or very near (within 10 cm) the ocean surface will be dead after 2.5 h of exposure. Under solar spectral irradiance simulating a 20% decrease in ozone layer thickness, this time drops to 2.2 h. These are first-order estimates based upon irradiance taken at a time of day during which the values would be maximal. Nonetheless, they illustrate the relative changes in UV-B effects that will result from ozone layer depletions expected over the coming decades. It is also important to point out that variability in cloud cover, water quality, and vertical distribution and displacement within the mixed layer, can all have a greater effect on the flux of UV-B radiation to which C. finmarchicus eggs are exposed than will ozone layer depletion at these latitudes. Received: 2 March 1998 / Accepted: 18 December 1998     

Carbon requirements of a population of the estuarine copepod Eurytemora affinis

The production of algae in the Patuxent River estuary (Maryland, USA) was much less than the carbon requirements of a population of the calanoid copepod Eurytemora affinis. Detrital carbon was present in quantities that suggest turnover times of 8 to 83 days for this pool, assuming that all algal production was consumed by E. affinis. We hypothesize that E. affinis must consume detritus to meet a part of its requirements for energy.     

In situ grazing rate of the copepod population in the western subarctic North Pacific during spring

Time series sampling with a multi-layer plankton sampler was carried out in the western subarctic North Pacific during spring 1991. Neocalanus cristatus, N. flemingeri and Eucalanus bungii dominated and accounted for 88.5% of the copepod population in volume. Neocalanus spp. were distributed in the upper mixed layer, while E. bungii was mainly distributed between 120 and 300 m throughout the day and night. In contrast, Metridia pacifica, Pleuromamma scutullata and Gaetanus simplex showed clear diel vertical migration. Grazing activities were estimated simultaneously by gut fluorescence. Nocturnal grazing was observed for diel migrating species. Neocalanus spp. did not have a diel feeding rhythm and had relatively low gut fluorescence. E. bungii was considered to be dormant during the observation period. The estimated grazing rate of the copepod population on phytoplankton was 1.4 to 2.0% of the primary production while the metabolic requirement was 8.3 to 12.4% of the primary production. These facts suggest that the copepod population was unimportant as primary consumers and that microzooplankton plays a much more important role in sustaining low standing stock of phytoplankton and a high nutrient concentration in the western subarctic Pacific Ocean.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Trophic specialisation of metazoan meiofauna at the Håkon Mosby Mud Volcano: fatty acid biomarker isotope evidence

We report the results of a detailed investigation on the trophoecology of two dominant meiofaunal species at the Håkon Mosby Mud Volcano (HMMV), a deep-sea cold methane-venting seep. Analyses of fatty acids (FAs) and their stable carbon isotopes were used to determine the importance of chemosynthetic nutritional pathways for the dominant copepod species (morphologically very similar to Tisbe wilsoni) inhabiting the volcano’s centre and the abundant nematode Halomonhystera disjuncta from the surrounding microbial mats. The strong dominance of bacterial biomarkers (16:1ω7c, 18:1ω7c and 16:1ω8c) coupled with their individual light carbon isotopes signatures (δ¹³C ranging from ?52 to ?81‰) and the lack of symbiotic relationships with prokaryotes (as revealed by molecular analyses and fluorescent in situ hybridisation) indicated that chemosynthetically derived carbon constitutes the main diet of both species. However, the copepod showed a stronger reliance on the utilisation of methanotrophic bacteria and contained polyunsaturated FAs of bacterial origin (20:5ω3 and 22:6ω3 with isotope signatures δ¹³C < ?80‰). Instead, the FA profiles of H. disjuncta suggested that sulphide-oxidising bacteria constituted the main diet of this nematode. Therefore, HMMV can be regarded as a persistent deep-sea cold seep, allowing a chemosynthesis-based trophic specialisation by the dominant meiofaunal species inhabiting its sediments. The present investigation, through the determination of the fatty acid profiles, provides the first evidence for trophic specialisation of meiofauna associated with sub-habitats within a cold seep.     

Evolution d'une population de copépodes dans le système des courants équatoriaux de l'Océan Pacifique. Zoogéographie, écologie et diversité spécifique

During the cruise “Alizé” along the Equator in the Pacific Ocean from 86° 20 W to 151° 01 E (Fig.1), 33 night tows were made with a 1 m plankton net, in the 0 to 200 m layer. The samplings show, 3 types of copepod distribution (Figs. 2 to 5); the main type presents the well-documented decrease from east to west. The different distributions seem controlled by: suface phytoplankton abundance in the case of Eucalanus sublenuis temperature and the thermocline in Euchaeta media and Pleuromamma quadrungulata; type and availability of prey in Euchaeta marina; upwelling and the Peru current in Eucalanus elongatus hyalinus and Rhincalanus nasutus. The 3 warm-water species Rhincalanus cornutus, Eucalanus attenuata and Eucalanus subcrassus are more abundant in the western part of the Pacific Ocean, where temperature is higher. The 2 cosmopolitan species Pleuromamma xiphias and P. abdominalis do not show any clear relationship with the biotope. Probably the presence of some species in the upper water layer in the zone of intense divergence and upwelling emphasizes the westward decrease. It has been possible to define 4 zones by phytoplankton distribution, currents, vertical circulation, and the physical and chemical properties of the biotope; these zones agree with the zonation of the copepods in accordance with the results of a rank test. Within these limits, the 4 indices of specific diversity I=-Σ pi log₂ pi and the equitibility have been calculated. The diversity variations account for the ecosystem evolution after Margalef's theory. The lower specific diversity in the oriental zone is related to the well developed divergence, the high rates of nutrients, the intense phytoplankton production, the instability in time and the southern water inflow; in this zone, the ecosystem is in an early stage of succession with a percentage of herbivores such as Eucalanus subtenuis (45%) present. The diversity index confirms the changes introduced by the trade winds along the Equator, mainly in the eastern part of the Pacific Ocean.     

Calculation and use of selectivity coefficients of feeding: Zooplankton grazing

A straightforward method for calculating selectivity coefficients (W_ij) of predation from raw data, mortality rates of prey, filtering rates, feeding rates and electivity indices is derived. Results from a comparison of selectivity coefficients for the copepod Diaptomus oregonensis grazing under a number of experimental conditions suggest that W_ij's for size-selective feeding are invariant, a conclusion also supported by the leaky-sieve model. Recommendations are made on how to use W_ij's in linear and nonlinear feeding constructs for zooplankton and other animals.     

Summer zooplankton distribution in a Georgia Estuary

Zooplankton were collected with Van Dorn bottles at two estuarine stations near Savannah, Georgia, USA, during summer, 1974, from the surface, 2 m, 4 m, and at the bottom of the water column at 2-h intervals for 11 days. Acartia tonsa occurred at the surface at night and during high and rising tides in higher concentrations than at the same depth during daylight and other tide stages. The highest bottom concentrations of this calanoid copepod occurred at high tide. A negative phototaxic response was also apparent in this species. While not quantitatively sampled by Van Dorn bottles, harpacticoid copepods and Pseudodiaptomus coronatus Were distributed in a manner similar to that of A. tonsa. Copepod nauplii, on the other hand, were always more abundant at the surface than at the bottom, with the highest concentration occurring during low tide. The distribution pattern of nauplii appears to be a function of dilution and concentration by tidally induced changes in water volume, whereas more advanced copepod stages follow a complexly controlled distribution pattern linked with tidal mixing dynamics as outlined in a previous study by Jacobs (1968).     

Analysis of phytoplankton pigments by HPLC before,during and after mass occurrence of the microflagellate Corymbellus aureus during the spring bloom in the open northern North Sea in 1983

Daily observations in April and May 1983 near a subsurface drifter launched in the Fladen Grounds area of the North Sea revealed that a large crop of the microflagellate Corymbellus aureus Green, producing up to 3 g C m^-2 d^-1, succeeded the diatoms dominating during the earlier phase of the spring bloom. The concentration of fucoxanthin was highest during the first half of May, while pigment fingerprints of high-performance liquid chromatograms of suspended matter sampled during the second half of May were dominated by 19-hexanoyloxyfucoxanthin, the main carotenoid of C. aureus (0.8x10^-9 mg cell^-1). This colonial Prymnesiophycean was described for the first time less than 10 years ago and has nerver been reported to be present in the North Sea. After 16 May the C. aureus population rapidly lost its growth potential, but the peak of the bloom in terms of cell number was found several days later (up to 9x10⁶ cells per dm³ on 19 May). The population then started to decline; senescence was associated with decreasing pigment contents of cells. The low growth rate of copepods registered during the second half of May was probably related to the poor quality of C. aureus as copepod food: the concentration of a phaeophorbide a typically found in copepod fecal pellets was highest during the diatom phase of the spring bloom preceding the C. aureus bloom.     

Behavioral adaptations of the cubozoan medusa <Emphasis Type="Italic">Tripedalia cystophora</Emphasis> for feeding on copepod (<Emphasis Type="Italic">Dioithona oculata</Emphasis>) swarms

The cubozoan medusa Tripedalia cystophora preys on dense swarms of the copepod Dioithona oculata in the mangrove prop-root habitat of Puerto Rico. The copepod swarms form in shafts of sunlight that penetrate the mangrove canopy during the day. T. cystophora are found primarily within the same illuminated areas, feeding heavily on the dense swarms of copepods. Laboratory studies were conducted to examine the behavioral adaptations of T. cystophora that allow them to remain within these dense copepod swarms. In the presence of vertical light shafts, T. cystophora showed both a significant change in swimming speed and a significant increase in turning rate. T. cystophora turn abruptly when they leave a vertical shaft of light and quickly re-enter. The presence or absence of copepod prey, either in swarms or dispersed, had no significant effect on T. cystophora swimming speed or turning behavior. Feeding rates of T. cystophora on the copepod D. oculata increase rapidly with copepod density, and maximum rates are not achieved until the high densities associated with swarms are reached. It seems doubtful that these small medusae could survive without their ability to encounter and exploit these dense swarms of copepods.     

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.