PHOSPHOGLUCOMUTASE ALLOZYME EVIDENCE FOR AN OUTCROSSING MODE OF REPRODUCTION IN THE HERMAPHRODITIC BROODING BIVALVE MYSELLA TUMIDA (GALEOMMATACEA)

Allozyme variation of a Mysella tumida population in PatriciaBay, B.C., Canada was investigated at the phosphoglucomutase(PGM) locus. Seven alleles were detected. The observed frequencyof allelic combinations did not differ significantly from randommating expectations (0.25 < P < 0.50), however, heterozygositylevels were slightly greater than expected (Selander's D = 0.022).These results, together with previous work on the M. tumidareproductive cycle, indicate that natural populations of thisbivalve rarely, if ever, self-fertilize. ^*Present Address: Friday Harbor Laboratories, University ofWashington, Friday Harbor, WA 98250, U.S.A. ^**Present Address: Museum of Zoology, University of Michigan,Ann Arbor, M1 48109, U.S.A. (Received 30 December 1986;     

Effects of diffusion and upwelling on the formation of red tides

In this paper, records on the timing and location of specificred tides monitored once or twice a week in Mikawa Bay, Japan,are related to horizontal and vertical mixing rates determinedfrom a numerical model. Horizontal (K_h) and vertical (K_z) diffusioncoefficients, and upwelling velocities, were estimated usinga box model analysis. In the wind-mixed period and in the upperlayer during the stratified period, K_h was estimated to be ofthe order of 10² m² s^–1. During the stratified period,K_z was estimated to be of the order of 10^–5 m² s^–1.The upwelling velocity was calculated to be in the range 0.35–5.1m day^–1 with an average of 1.5 m day^–1. Comparisonbetween the literature values of the specific growth rate (µ)of the red tide-forming diatoms and calculated K_h values duringthe red tides show that diatoms which have a low µ cannotform red tides in a strongly diffusive environment, while specieshaving a high µ can form red tides even in a strong diffusiveenvironment. On the other hand, no clear relationship was foundbetween µ of the flagellate group and K_h, although theflagellate group formed red tides even in severe diffusive conditions.From the comparison between the literature values of sinkingrate and swimming speed and the physical parameters associatedwith vertical processes, it was concluded that flagellates willform red tides, even in severe diffusive conditions, by usingtheir swimming ability, while diatoms form red tides by theirhigh growth rates with the aid of vertical diffusion and theupwelling movement of water.     

Hydrodynamic disturbances produced by small zooplankton: case study for the veliger larva of a bivalve mollusc

Zooplankton produce hydrodynamic disturbances during swimmingand feeding that enlarge their perceptive volume. From the standpointof both prey and predators, fluid disturbances increase theprobability that an organism is detected, identified and reactedto within appropriate time and space scales. Morphology andkinematics dictate the magnitude, symmetry and attenuation ofdisturbances in the fluid medium. Therefore, fluid disturbancesmay be species and age (size) specific. Normal and high-speedvideo microscopy was used to study flow-field generation byfree-swimming and tethered bivalve larvae. These organisms swimand feed using many highly coordinated and symmetrically distributedappendages (i.e. cilia). Larvae tethered in flow at variousfree stream velocities (U₀), simulating swimming activity, inducedparticle trajectories approximately parallel to the organism'sdorso-ventral axis. Velocity (v) and acceleration (a) were symmetricalin the transverse plane and asymmetrical in the vertical plane.Greatest velocity magnitudes ({small tilde}7, 3 and 6 mm s^–)occurred dorsal to the velum and attenuated with source distance(r) as 1/r, 1/r^1.9 and 1/r^2.9 at 10 s^– U₀ =, 3.1 and 6.4mm s^–1, respectively. For a larva in flow, but with velumretracted, simulating sinking, velocity attenuated at Mr towardsthe organism. Mean velocity gradients were on the order of 3,8 and 10 s^–1 for swimming, sinking and hovering larvae,respectively. The high-frequency (22 Hz) component of particlevelocity past free-swimming larvae was due to beat frequencyof the velar cilia. This attenuated rapidly with r leaving onlylow-frequency (1–3 Hz) disturbances 0.1 mm beyond thetips of the cilia. Comparisons of the kinetic energy dissipationrate for turbulence in coastal waters with the kinetic energyof laminar flow fields implied possible dominance of the flowfield of hovering, but not swimming, larvae to at least threebody diameters from the organism (–1 mm). These differencesin flow fields have important implications for larval survival.The perceptive volume of a hovering larva will be 40-fold greaterthan that of a swimming or sinking larva. However, a hoveringlarva U also more likely to be detected by a potential predatorthat uses mechanosensory organs to locate prey.     

The Development of Fast-Start Performance in Fishes: Escape Kinematics of the Chinook Salmon (Oncorhynchus tshawytscha)

C-starts are high acceleration swimming movements critical forpredator avoidance by fishes. Since larval fishes are particularlyvulnerable to predation, C-start behavior is likely to be especiallyimportant during early life history stages. This paper examinesthe developmental changes in C-start performance with kinematicdata on immature chinook salmon (Oncorhynchus tshawytscha) (eleuthroembryostage, sensu Balon, 1975). The scaling of C-start kinematicsof immature fishes differs from that of adults. Adult C-startdurations increase with increasing body length while C-startdurations of immature fishes decrease (e.g., adult stage 1 duration[sec] = 0.0019.length [L] [cm] $ 0.026 [R² = 0.77] [Webb, 1978];eleuthroembryos stage 1 duration [sec] = –0.026L [cm]$ 0.100 [R² = 0.81]). Distance traveled during stage 2 alsodiffers between adult and immature fishes. Adult distance traveledscales directly with length (distance [cm] = 0.38L^1.01 [cm],R² = 0.96 [Webb, 1978]) while chinook eleuthroembryo distancetraveled is positively allometric with length (distance [cm]=0.37L¹³¹ [cm], R² = 0.83). There are similarities in the developmentof C-starts and burst swimming. For example, mean velocity scalessimilarly between the two locomotor modes (For burst swimming:U_mean [cm/sec] = 8.1 ± 1.1L [cm] $ 4.89 [R² = 0.86] [Webband Corolla, 1981]. For C-start stage 2: U_mean [cm/sec] = 10.96L[cm] - 14.09 [R² = 0.70]). This study demonstrates that C-startescape performance improves during early post-hatching development.Comparisons of immature chinook salmon fast-starts with dataon larval burst swimming and on adult C-starts suggest thatchanges specific to developing fish affect the scaling of kinematicparameters.     

Identification of sibling species within the “Sarsia tubulosa complex” at Friday Harbor,Washington (Hydrozoa: Anthomedusae)

The anthomedusa Sarsia tubulosa has long been considered a “variable” species at the Friday Harbor Laboratories and elsewhere. A series of morphological and behavioral observations, coupled with an extensive program of hybridization testing, has revealed the existence of at least three species of large Sarsia at Friday Harbor: Sarsia tubulosa (M. Sars, 1835), Sarsia princeps (Haeckel, 1879), and an undescribed species. Sarsia tubulosa and the undescribed species make up the bulk of the population. S. princeps is rare. The common species hybridize at such a low level in the laboratory that hybrids in nature are probably rare. Potential isolating mechanisms consist of a one-hour time difference in spawning (in the laboratory) and a partial species specificity of sperm chemotaxis. Since manipulation of the spawning time does not affect the level of hybridization and the species specificity of sperm chemotaxis is incomplete, some other mechanism of reproductive isolation is probably operating to maintain species separation in nature. The hybridization experiments do reveal a relationship between the level of species specificity of sperm attraction and the amount of hybridization, suggesting that sperm chemotaxis plays a role in fertilization in these animals.     

Light, temperature and nitrogen as interacting factors affecting diel vertical migrations of dinoflagellates in culture

Diel vertical migrations of the marine dinoflagellates Gonyaulaxpolyedra Stein and Ceratium furca (Ehr.) Clap, et Lachm. werefollowed in a laboratory tube (2.02 m x 0.25 m) under a 12:12hlight:dark cycle. The effects of temperature stratification,two levels of surface irradiance and nitrogen depletion on patternsof vertical migrations were examined. At temperatures between22–26°C with small temperature gradients, both speciesmigrated at a rate of 0.7 –1.0 m h^–1. Steeper thermoclines(ca. 0.8°C 0.1 m^–1) with temperatures below ca. 20°Ccaused a marked decrease in swimming speed which resulted inaccumulations of cells in these thermocline regions. Under conditionsof nutrient sufficiency both algae migrated into the surfacelayers at irradiance values of over 1000 µE m^–2s^–1. Increasing nitrogen depletion caused the downwardmigration of both algae to commence progressively earlier inthe day and before the end of the light period. The earlierdownward migrations enabled a more complete descent throughthe thermocline. Nitrogen depleted cells of Gonyaulax continuedto undertake vertical migrations but avoided high irradiancesthus forming subsurface maxima at irradiance levels close to150 µE m^–2 s^–1. Ceratium cells which exhaustedboth inorganic nitrogen and phosphorus ceased to migrate accompaniedby a large change in cellular fluorescence.     

Sublethal effects of the toxic dinoflagellate Karenia brevis on marine copepod behavior

Apart from grazing interactions, little is known regarding thesublethal effects of Karenia brevis cells on copepod behavior.We conducted grazing and mortality experiments with K. breviscells and brevetoxins (PbTx-2), establishing routes of toxicityfor the copepods Acartia tonsa, Temora turbinata and Centropagestypicus. Subsequent behavioral experiments determined whethercopepod swimming and photobehavior, both behaviors involvedin predator avoidance, were impaired at sublethal K. brevisand PbTx-2 levels. Copepods variably grazed toxic K. brevisand non-toxic Prorocentrum minimum at bloom concentrations.Although copepods accumulated brevetoxins, significant mortalitywas only observed in T. turbinata at the highest test concentration(1 x 10⁷ K. brevis cells L^–1). Acartia tonsa exhibitedminimal sublethal behavioral effects. However, there were significanteffects on the swimming and photobehavior of T. turbinata andC. typicus at the lowest sublethal concentrations tested (0.15µg PbTx-2 L^–1, 1 x 10⁵ K. brevis cells L^–1).Although physiological incapacitation may have altered copepodbehavior, starvation likely played a major role as well. Thesedata suggest that sublethal effects of K. brevis and brevetoxinon copepod behavior occur and predicting the role of zooplanktongrazers in trophic transfer of algal toxins requires knowledgeof species-specific sublethal effects.     

The use of underwater camera in studies of vertical distribution and swimming behaviour of a sergestid shrimp, Sergia lucens

The vertical distribution and swimming behaviour of Sergia lucenswere investigated by means of underwater photography. Duringthe day the species was abundant from 3 to 7 m above the bottomon the upper continental slope at depths from 210 to 360 m.No shrimp were observed closer than 0.5 m above the bottom.The maximum concentration observed was 6.1 individuals m^–3.It is speculated that in many areas on the continental slopesand rises, the density of zooplankton and micronekton is muchhigher near the bottom in the day than in the overlying midwater,and that this may provide a concentrated pelagic food for predatorsclose to the seafloor.     

Vertical distribution of pelagic chaetognaths and feeding of Sagitta enflata in the Central Equatorial Pacific

The vertical distribution and feeding of pelagic chaetognathsat 5°S, 160°W in the Central Equatorial Pacific wereinvestigated using a series of 0–500 m vertical haulswith a VMPS net over a 24 h period between 6 and 7 October 1990.The total number of individuals per haul was between 370 and688. Fourteen species in four genera were found at this station.The most abundant species was Sagitta enflata which comprised32.4–61.1% of the individuals collected from the 0–500m layer. Mesopelagic species made up 9.3–15.1% of thetotal number of individuals. Sagitta enflata and Pterosagittadraco were found in the upper part of the thermocline both byday and at night. The fraction of the population containingfood items (FCF) of S.enflata in the 0–50 m layer variedbetween 4.8 and 12.5% (mean 10.8%) and feeding activity washighest between sunrise and noon. The percentages of Copepoda,Foraminifera, crustacean larvae, Chaetognatha, Pteropoda, Ostracoda,fish and unidentified material in the gut of S.enflata were51.9,6.7,3.8,2.9,1.9,1.9 and 30.9%, respectively. Sagitta enflataconsumed food organisms which were mainly between 0.5 and 1.0mm in length. The daily feeding rate of S.enflata was 1.81 preyper individual, which was equivalent to 8.06 mg C m^–2day^–1. This corresponded to     

Infection of Coscinodiscus spp. by the parasitoid nanoflagellate Pirsonla diadema: I. Behavioural studies on the infection process

The investigation of successive steps involved in the infectionprocess of the marine diatoms Coscinodiscus granii and Coscuwdiscuswailesii by the host-specific parasitoid nanoflagellate (PNF)Pirsonia diadenw showed that flagellates reacted chemokJinokineticallywith changes of swimming pattern to the presence of a host diatom.Chemosensory stimulation appeared to induce readiness for infection,whereas attachment and penetration of the diatom cell wall wasinduced by a mechanosensory response to morphological featureson the diatom frustules. The mean swimming speed of P.diademaflagellates decreased during their infective lifetime of 3 daysfrom an average of 78 µm s^–1 to 51 µm s^–1while the frequency of small loops in the swimming pattern increasedfrom 0.8 to 6.3 loops min^–1. At high Cgranii densities,an epidemic was delayed. It is suggested that this could becaused by overlapping gradients of extracellular material releasedby the diatoms which impaired the sensing of spatial gradientsby PNF and, therefore, the location of hosts.     

Size-specific swimming behavior of Daphnia pulex

Body length affects several aspects of the behavior of quietlyswimming Daphnia pulex Swimming and sinking rates were measuredat 0.033 s intervals during the ‘hops’ characteristicof Daphnia swimming behavior Larger animals swim faster, covermore distance, and produce more powerful swimming strokes. LargerDaphnia also sink faster, but the sinking rate scales as lengthto the 0 58 power, far lower than the power of 2 00 predictedby Stokes Law considerations. The number of hops s^–1 wasindependent of body size, although a theoretical analysis predictshopping rate (antennal beat frequency) should increase as bodylength squared. Turning behavior, measured as the ratio of displacementto total distance, during 5 s, is also independent of body sizeIndependence of several parameters of body motion and body sizeimplies that factors other than simple mechanics affect Daphniaswimming behavior     

The effects of food deprivation, prey density and volume on clearance rates and ingestion rates of Diacyclops thomasi

The ingestion rates of the copepod, Diacyclops thomasi, on thesoft-bodied rotifer, Synchaeta pectinata, increased 10-fold(0.07–0.77 Synchaeta h^–1) over the range 50–250prey l^–1. The saturating functional response curve appearedsigmoid but was statistically indistinguishable from a parabola.The response curve was more linear and 10 times lower over thesame range of density when Diacyclops was offered Kerarellacochleans, a species having a stiffened lorica. Diacyclops maximizedits ingestion rate on Synchaeta as a function of the availablegut space. Predation effort, measured as clearance rates, waslinked tentatively to changes in swimming speed of Diacyclopsand was a function of hunger level. Diacyclops, which were starvedfor varying periods of time, increased their ingestion rateson Synchaeta up to a maximum (-3.0 h^–1) after 7–10h of food deprivation. The gut passage time of Diacyclops wasestimated to be 7–8 h. Therefore, ingestion rates (andclearance rates) appeared to be strongly correlated to the volumeof food in the gut.     

An empirical model of primary productivity (14C) using mesocosm data along a nutrient gradient

The two parameters of the hyperbolic tangent equation, P_m and, were estimated from in situ vertical profiles of primary productionusing mesocosm data along a nutrient gradient. The parameters,derived from 4-h (around noon) ¹⁴C incubations, were used togetherwith the photosynthesis-light curve and hourly solar radiationdata to calculate daily primary production rates (P_d). Approximately40% of the daily production occurred in the 4 h around noon.Considering parameter uncertainty, there was no indication ofan increase in variation in production with increased nutrientloading, nor did biomass-specific P-I parameters increase. Annualproduction ranged from 82 to 901 g C m^–2 year^–1and was highest in the highest nutrient treatment tank. Dailyproductivity ranged from 0.02 to 9.1 g C m^–2 day^–1and was significantly correlated, in all treatments, with acomposite parameter BI₀/k (where B is phytoplankton biomass;I₀ is daily radiation and k is the extinction coefficient).Linear regressions of P_d against BI₀/k indicated that much ofthe variability (86%) in productivity was explained by lightavailability and phytoplankton biomass. Two approaches for predictingproductivity were compared: (i) predicting production directlyfrom environmental variables (i.e. BI₀/k) and (ii) predictingthe parameters of the P-I curve from environmental variablesand using these to calculate daily production.     

FEEDING AND GROWTH OF THE AEOLID NUDIBRANCH CUTHONA NANA (ALDER AND HANCOCK, 1842)

Feeding and growth of the aeolid nudibranch Cuthona nana wasdocumented. Cuthona nana feeds specifically on the colonialhydroid Hydractinia echinata. Nudi-branchs were observed onhermit crab shells in Gos-port Harbor at the Isles of Shoals,Maine and on old bridge pilings near Gerrish Island, Maine. Grazing rates of nudibranchs (3–25 mm) were quantifiedover 24 hour periods in the laboratory. Larger nudibranchs consumedmore hydroid tissue feeding mostly on polyps, whereas smallerindividuals consumed less and fed on mat tissue. Grazing intensityin the field at Gosport Harbor was determined by estimatingthe area of grazed patches of hydroid colonies attached to hermitcrab shells. The average patch size was 5.75% of the total colonyarea. Growth rates for nudibranchs were determined in the laboratory.Growth rates were greatest prior to maturation and oviposition.Animals from Gerrish Island and Gosport Harbor decreased insize after oviposition due to senescence. Nudibranchs from GerrishIsland reached a maximum size of 12 mm, compared to 23 mm foranimals at Gosport Harbor. ^*Present address: Division of Science and Mathematics. Universityof Minnesota, Morris. Minnesota 56267, USA     

The role of feeding behavior in sustaining copepod populations in the tropical ocean

A fundamental question regarding marine copepods is how themany species coexist and persist in the oligotrophic environment(i.e. Hutchinson’s paradox). This question is addressedwith a stochastic, object-oriented Lagrangian model that explicitlysimulates the distinct foraging behaviors of three prominenttropical species: Clausocalanus furcatus, Paracalanus aculeatusand Oithona plumifera. The model also individually tracks allprey cells. Each particle’s motion combines sinking, turbulentdiffusion and active swimming when applicable. The model successfullysimulates observed size-partitioned carbon uptake rates. Basedon the model results, the wide-ranging translational ambit employedby C. furcatus is best suited for the acquisition of passiveprey while the relatively stationary behavior of O. plumiferapromotes the capture of larger, quickly sinking cells. The modelresults further suggest that the slow velocities and feedingcurrent employed by P. aculeatus are best suited for acquiringthe smallest cells though it also has a slight advantage overC. furcatus in acquiring the largest prey. A resource threshold,at a prey concentration of 530 cells mL^–1, is consistentlyexhibited by all three modeled species. Overall, these resultsimply that the size-partition preferences due to their differentforaging behavior contribute to the coexistence of these threespecies. This paper is one of six on the subject of the role of zooplanktonpredator–prey interactions in structuring plankton communities.     

Egg production and hatching success in the peri-Antarctic copepod Calanus simillimus

In situ rates of egg production and hatching success are reportedfor Calanus simillimus, one of the most abundant calanoid speciesin peri-Antarctic regions, during the Italian ‘Italics’cruise in the Straits of Magellan in March-April 1995. Low fecundity(8.6 eggs female^–1 day^–1) and fecal pellet production(0.8 fecal pellets female^–1 day^–1) in this periodindicate that the species was feeding very poorly. Sixty-sixof the 126 females sampled did not produce eggs and 80 of thesedid not produce any fecal pellets during the 24 h period ofincubation. Striking abnormal naupliar and embryonic developmentwas recorded in 81.8% of the eggs spawned. Aberrant eggs didnot undergo normal cleavage, and failed to develop to hatching.Deformed nauplii were asymmetrical and presented strong anatomicalanomalies in the total body length and number of swimming appendages.These results are discussed in the light of recent findingson the causes of low hatching success in copepods.     

Temporal Changes in Swimming Direction during the Phototactic Orientation of Individual Cells in Cryptomonas sp.

The response of individual Cryptomonas cells to continuous lightwas recorded using infrared video-micrography. Swimming directionsand temporal shifts in swimming direction of each cell weremeasured. White light of 0.1–1 W m^–2 elicited apositive phototactic orientation, but did not induce any photophobicresponse. Light of 100 W m^–2 induced a photophobic responseat the onset of actinic irradiation, but did not induce positivephototactic orientation. No correlation between positive phototacticorientation and photophobic response was found in this species.The direction toward the light source was defined as 0°,and the direction away from the source as 180°. Within 2s after the onset of lateral monochromatic light of 570 nm at0.1 W m^–2, cells which were swimming in a direction ofless than 120° predominantly shifted their course towardthe light source. Cells swimming in directions of larger than120° shifted their course as randomly as those in the dark.Thus, for phototactic orientation, the cells must perceive thelight from their anterior side. (Received July 29, 1985; Accepted November 4, 1985)     

The scaling and structure of aquatic animal wakes

Animal generated water movements are visualized and quantifiedusing two-dimensional particle image velocimetry (PIV). Theresulting vector flow fields allow for the study of the distributionof velocity, vorticity and vortices. Structural and temporalaspects of animal-induced flows covering a range of Reynolds(Re) numbers between less than 1 to more than 10⁴ are presented. Maps of flow induced by continuous foraging and intermittentescape responses of tethered nauplius and copepodid stages ofthe marine copepod Temora longicornis offer insight in viscosity-dominatedflow regimes. Fast escape responses of the equally sized largestnauplius stage and the smallest copepodid stage are compared.The nauplius moves by generating a viscous flow pattern withhigh velocities and vorticity; the copepodid moves by usinginertial effects to produce a vortex ring with a rearward jetthrough the center. Larvae and small adult fish (zebra danio) use a burst-and-coast-swimmingmode at Re numbers up to 6,000, shedding a vortex ring withthe associated jet at the tail during the burst phase. Flowpatterns during the coasting phase differ between the smalllarvae and larger adults due to the changes in importance ofviscosity. A 12 cm long mullet swimming in a continuous mode generatesa chain of vortex rings with a backward undulating jet throughthe centers of the rings at Re numbers of 4 x 10⁴ in inertia-dominatedregimes. Our empirical results provide realistic insight in the scaleeffects determining the morphology of the interactions betweenanimals and water.     

Vertical distributions, diel and ontogenetic vertical migrations and net avoidance of leptocephali of Anguilla and other common species in the Sargasso Sea

Day- and night-time vertical distributions and their ontogeneticchanges in Anguilla leptocephali and other common species ofleptocephali were determined and compared during five cruisesin the Sargasso Sea using an opening - closing 2-m ring netto sample discrete depth strata between 0 m and 350 m deep.No difference in vertical distribution was ever found betweenAnguilla rostrata (American eel) and A. anguilla (European eel).Anguilla leptocephali <5 mm long did not exhibit a diel verticalmigration, as they were distributed between 50 m and 300 m bothby day and by night. The vertical distribution of these smalllepto-cephali is probably roughly representative of the depthdistribution of adult spawning. Anguilla     

Ecological Studies of Seaweeds in McMurdo Sound, Antarctica

Three species of benthic marine macroalgae comprise the chiefcomponents of the seaweed flora of McMurdo Sound, Antarctica.Quantitative studies at Cape Evans demonstrate a depth-relateddistribution pattern, with Iridaea cordata (Turner) Bory inshallow water, Phyllophora antarctica A. and E. S. Gepp abundantand fertile at intermediate depths, and Leptophytum coulmanicum(Foslie) Adey dominant below 20 m. The vertical distributionof species is correlated with irradiance levels. At sites withthinner annual sea ice and less snow accumulation (e.g., CapeEvans, Cape Royds, and Granite Harbor), vertical distributionsare shifted downward relative to those at sites that remaincovered most of the year with thick or snow-covered fast ice(e.g., Cape Armitage and New Harbor). Disturbance caused byice scour and anchor ice probably determines the upper limitof algal distribution; herbivory is apparently absent. There is a disproportionate representation of cystocarpic femalegametophytes in populations of I. cordata and P. antarctica.Perennation via persistent basal crusts and apogamic recyclingof gametophytes are suggested as factors structuring phase distribution.Although the overall size of I. cordata blades is greater (andcystocarpic plants more abundant) at Cape Royds than at CapeEvans, cystocarpic blades are similar in minimum size at bothsites, indicating a threshhold size or age for the onset ofreproduction in this species.