Multifarious activities of gut epithelium in an appendicularian (Oikopleura dioica: Tunicata)

The functions of the various gut regions of Oikopleura dioica (oesophagus, left and right gastric lobes, vertical intestine, mid-intestine and rectum) were investigated by means of histochemical, histoenzymatic and immunohistochemical techniques at light and electron microscopes. Ciliary food progression is evidenced by the presence of ATPases on cilia, along the entire gut, with the exception of the cardiac valve, a passive device controlling food direction. Absorptive processes (alkaline phosphatase), active transport (ATPases) and nitrogen excretion (D-amino acid oxidase) occur along the entire gut, in both ciliated microvillar and globular cells. The latter, typical of the left gastric lobe and rectum, are also involved in endocytotic processes (exogenous peroxidase as tracer) and intracellular digestion (!-amylase, aminopeptidase M, acid phosphatase, 5'-nucleotidase, non-specific esterase). The giant cells of the gastric band participate in extracellular digestion; they contain secretory granules positive to various hydrolytic enzymes, the activity of which is also recognisable in faecal pellets inside the intestinal lumen. Lipid storage occurs mainly in the right gastric lobe and vertical intestine, whereas protein storage takes place in the rectal granular cells. Epithelial transport and possible osmoregulation occur along the entire gut, especially at the level of diffuse baso-lateral interdigitations, which increase the plasmalemma surface enormously, are often associated with mitochondria and possess numerous ATPase pumps. Data extend previous histological observations and hypotheses on the physiological role of the various gut regions. The remarkable and specific location of enzymatic activities and nutrient storage are in agreement with the high capacity of O. dioica to process a great quantity of food very rapidly and efficiently.     

Seasonal variations in the densities of fecal pellets produced by Oikopleura vanhoeffeni (C. Larvacea) and Calanus finmarchicus (C. Copepoda)

Two abundant macrozooplankters, Oikopleura vanhoeffeni (Lohmann) and Calanus finmarchicus (Gunnerus) were collected from the coastal waters off Newfoundland in different seasons during 1990–1991 and incubated in natural seawater to collect freshly egested, field produced, fecal pellets. The densities of fecal pellets from O. vanhoeffeni and C. finmarchicus were measured in an isosmotic density gradient. These are the first reported seasonal measurements of fecal pellet densities from two different types of macrozooplankters, O. vanhoeffeni, a larvacean, filter feeder and C. finmarchicus, a crustacean, suspension feeder. Pellet density ranges and medians were significantly different among seasons for both species, depending primarily on the type of phytoplankton ingested and its ability to be compacted. Winter O. vanhoeffeni and fall C. finmarchicus feces filled with nanoplankters and soft bodied organisms had less open space [packing index (% open area) = 3.5 and 4% for O. vanhoeffeni and C. finmarchicus, respectively] and were more dense (1.23 and 1.19 g cm^-3) than spring feces filled with diatoms (packing index = 15 and 23%, density = 1.13 and 1.11 gcm^-3). For copepods, these results contrast with previously published density values and with the predicted copepod fecal pellet density calculated, in the present study, using the conventional mass/volume relationship. Copepod spring and summer diatom-filled feces had a calculated density of 1.12 and 1.24 gcm^-3 vs a measured median density of 1.11 gcm^-3 for both spring and summer feces; the fall feces containing nanoplankters had a calculated density of 1.08 gcm^-3 vs a measured median density of 1.19 gcm^-3. Knowledge of the seasonal variations in fecal pellet densities is important for the development of flux models.     

Predator-prey interactions between the larvacean Oikopleura dioica and bacterioplankton in enclosed water columns

The larvacean Oikopleura dioica Fol was fed ³H-labeled natural assemblages of marine bacterioplankton. Grazing rates ranged from <1 to >100 ml day^-1 individual^-1 and were highly dependent onlarvacean body size. These rates were combined with estimates of abundance of O. dioica in large floating enclosures with semi-natural populations (Controlled Ecosystems Populations Experiment, CEPEX) to determine the impact of the larvacean on the bacterial populations and to estimate the amount of bacteria ingested by the larvaceans. Apparently, O. dioica has minimal influence on the population dynamics of the free-living bacteria, although bacteria may form a substantial portion of the larvacean's diet.     

Effects of ocean acidification, temperature and nutrient regimes on the appendicularian Oikopleura dioica: a mesocosm study

Increasing pCO₂ is hypothesized to induce shifts in plankton communities toward smaller cells, reduced carbon export rates and increased roles of gelatinous zooplankton. Appendicularians, among the most numerous pan-global “gelatinous” zooplankton, continuously produce filter-feeding houses, shortcutting marine food webs by ingesting submicron particles, and their discarded houses contribute significantly to carbon fluxes. We present a first mesocosm-scale study on the effects of temperature, pCO₂ and bloom structures on the appendicularian, Oikopleura dioica. There were effects of temperature and nutrients on phytoplankton communities. No shifts in functional phytoplankton groups, nor changes in particle sizes/morphotypes, known to impact appendicularian feeding, were observed under manipulated pCO₂ conditions. However, appendicularian abundance was positively correlated with increased pCO₂, temperature and nutrient levels, consistent with hypotheses concerning gelatinous zooplankton in future oceans. This suggests appendicularians will play more important roles in marine pelagic communities and vertical carbon transport under projected ocean acidification and elevated temperature scenarios.     

Field behavior and adaptive strategies of appendicularians (Chordata: Tunicata)

The behavior of 7 species of appendicularians from the family Oikopleuridae was observed using SCUBA in the Gulf of California and the Florida Current. The frequency and orientation of feeding and the pattern of swimming while within the house varied considerably among species. Appendicularians expanded new houses in 1 1/2 to 5 min. House-expansion behavior was complex and variable among species. Appendicularians rarely abandoned the house in response to predation. Predators included the sergeant major (Abudefduf sp.), medusae, chaetognaths and ctenophores. The abandonment of the house was based on a cost-benefit behavior strategy which maximized time spent inside the house while minimizing predation and energy investment in house secretion. Strategies of predator avoidance are discussed.     

Influence of temperature changes on oxygen uptake and ammonia and phosphate excretion,in relation to body size and weight,in Oikopleura dioica (Appendicularia)

The relationship between the rates of oxygen consumption, ammonia and phosphate excretion of a pelagic tunicate, the larvacean Oikopleura dioica Fol, 1872 were assessed as a function of size, dry weight and ash-free dry weight at 15°, 20° and 24°C. O. dioica has higher respiration and excretion rates than copepods of similar weight, but the weight exponent of the allometric power function: Y=aX ^b is similar to that of other poikilotherms. Temperatures above 20°C have a depressing effect on respiration and ammonia excretion. 90% of the variance in metabolic rates is explainable by body mass and temperatures Q₁₀ values for oxygen consumption, ammonia and phosphate excretion, respectively, are 2.45, 1.86 and 1.75 between 15° and 20°C, and 3.75, 2.90 and 3.60 between 20° and 24°C. Metabolic quotients (O:N, O:P, N:P) indicate a protein-oriented diet. The results of this study suggest weak metabolic regulation in O. dioica, an energetic strategy which allows an immediate response to favourable changes in feeding conditions.     

Timing of sperm shedding and release of aggregates in the salp Thalia democratica (Urochordata: Thaliacea)

Many holoplanktonic forms, such as medusae, spawn in response to environmental cues such as light or temperature. Few observations of the time of reproduction have been made in salps. Small groups of mature aggregates of the salp Thalia democratica (Forskal, 1775), collected off the Great Barrier Reef of Australia, were held under constant light in early to mid-December in the years 1990, 1993 and 1994. Sperm were shed between 0300 and 0600 hrs each day with a mean time of about 0450 hrs for all individuals for all years combined. Solitary stages released chains of fertilizable female individuals between 0500 and 0700 hrs each day, with a mean time of 0540 hrs. Field observations suggested that similar timing occurred in nature. Advancing the time of dusk from 1900 to 1700 or 1800 hrs. did not significantly alter the mean time of sperm shedding or chain release. T. democratica aggregates, collected in the Mediterranean Sea in 1995 and maintained in constant light, had a broader distribution of individual spawning times, with a mean time of 0625 hrs, which differed significantly from that of the Australian population. Coordination of the events of sexual reproduction in T. democratica may improve fertilization success and be responsible for recognizable age cohorts within blooms of this species. Received: 27 May 1997 / Accepted: 20 June 1997     

Genetic diversity and genetic structure in the brown alga Halidrys dioica (Fucales: Cystoseiraceae) in Southern California

Genetic diversity and genetic structure in a population of the brown seaweed Halidrys dioica Gardner were evaluated in five sites in southern California, USA, in 1991, using isozyme electrophoresis. H. dioica is relatively long-lived and has an outcrossing mating system and floating reproductive fronds with the potential for longdistance gamete dispersal. Because these characteristics are hypothetically important in determining genetic diversity and structure, we predicted that genetic diversity would be high and genetic structure would be exhibited only at relatively large geographic scales in H. dioica populations. The data were consistent with the prediction: genetic diversity (% polymorphic loci, no. of alleles/locus, average expected heterozygosity) was high compared to that of other seaweed species. Genetic structure (Wright's F statistics, Nei's genetic distance, point-pattern analysis of alleles) was low within and among distinct rocky reefs over 4 km of coast but high in subpopulations separated by 90 km. Life-history characteristics may be useful predictors of genetic diversity and structure in seaweed populations, but information on selection regimes, long-distance dispersal, and the extent of clonal propagation, for example, are critically lacking.     

Correlation of bioluminescence emissions with ventral photophores in the mesopelagic squidAbralia veranyi (Cephalopoda: Enoploteuthidae)

Specimens of the squidAbralia veranyi were collected off the Bahamas in 1989. The bioluminescence of the ventral photophores was recorded on videotape at 11 to 12°C and 24°C, in conjunction with measurements of the spectral emission at the same temperatures. At 11 °C, the spectrum was unimodal, peaking at 490 nm with a narrow bandwidth. At the higher temperature a shoulder at about 440 nm appeared in the emission spectrum. The short, bright flashes from the subocular photophores had a broader bandwidth and a shorter wavelength emission maximum (475 nm) than that of the ventral photophores. Video recordings at the two temperatures showed no changes in either the identities of the luminescing photophores or their relative intensities. One structurally distinct type of photophore was not illuminated. We conclude that the observed spectral changes were produced within one group of photophores, and that recruitment of the unilluminated photophores would produce an additional spectral component, previously reported from another species ofAbralia (A. trigonura).     

Trophodynamic function of copepods,appendicularians and protozooplankton in the late summer zooplankton community in the Skagerrak

The study was carried out in the Skagerrak during late summer when population development in the pelagic cycle culminated in the yearly maximum in zooplankton biomass. The cyclonic circulation of surface water masses created the characteristic dome-shaped pycnocline across the Skagerrak. The large dinoflagellate Ceratium furca dominated the phytoplankton biomass. Ciliates and heterotrophic dinoflagellates were the major grazers and, potentially, consumed 43–166% of daily primary production. The grazing impact of copepods was estimated from specific egg production rates and grazing experiments. The degree of herbivory differed between species (14–85%), but coprophagy (e.g. feeding on fecal pellets) and ingestion of microzooplankton were also important. The appendicularian Oikopleura dioica was present in lower numbers than copepods, but cleared a large volume of water. The grazing impact of copepods and O. dioica was estimated to 57±24% and 12±12% of daily primary production, respectively. Sedimentation of organic material (30 m) varied between 169 and 708 mg C m^–2 day^–1, and the contribution from the mesozooplankton (copepod fecal pellets and mucus houses with attached phytodetritus of O. dioica) was 5–33% of this sedimentation. Recycling of fecal pellets and mucus houses in the euphotic zone was 59% and 36%, respectively. However, there was a high respiration of organic material by microorganisms in the mid-water column, and 34% of the sedimenting material actually reached the benthic community in the deep, central part of the Skagerrak.     

Architecture of,and water circulation and flow rate in,the house of the planktonic tunicateOikopleura labradoriensis

The gelatinous house ofOikopleura labradoriensis (Tunicata, Appendicularia), collected from the docks of Friday Harbor Laboratories, University of Washington, USA, in 1984, 1986 and 1990, was examined in vivo by stereomicroscopy and strobe-light macrophotography, and after fixation and processing for light and electron microscopy. In addition to confirming previous knowledge and adding new information on structural organization of the oikopleurid house, this study presents quantitative data on important aspects of its function. Particles small enough to pass through the inlet filters (pore width ~13 µm) were concentrated between differently constructed upper and lower food-concentrating filters (pore widths 0.18 and 0.24 µm, respectively). These filtes were held together by an intermediary screen of widely separated ribbon-like filaments. Water sieved through the filters left the house through a pressure-regulated exit valve. However, the intermittent activity of the tail pump and the elasticity of the house caused frequent refluxes of water that cleared both inlet filters and foodconcentrating filter screens of adhering particles. During these refluxes the food-concentrating filters usually collapsed and compacted the trapped particles into coarser aggregates. With each pumping cycle the particles and aggregates were brought closer to the midline. From here they were periodically drained into the mouth of the organism through a medial food-collecting tube, to be recaptured in a pharyngeal feeding filter secreted by the organism's endostyle. Based on the size and movements of the tail within the close-fitting tail chamber, a water flow rate of ~0.84 ml min^–1 was calculated for medium-sized houses (belonging to individuals with trunk length of ca. 1.2 mm). Taking the intermittent pumping activity of the tail into account, this equals ~35 ml h^–1. Flow through the food-collecting tube was ~1 µl min^–1, laminar and intermittent, and was probably comparable to a rate of ~0.04 ml h^–1. Accordingly, the house allowed the oikopleurid to feed on a ca. 1000 × concentrated suspension of particles. Water speed through the meshes of the food-concentrating filters was ca. 0.15 mm min^–1, or 2.5 µm s^–1.     

Measurement of in situ clearance rates of Oikopleura vanhoeffeni (Appendicularia: Tunicata) from tail beat frequency, time spent feeding and individual body size

In situ observations (1994 to 1996) of the behavior of Oikopleura vanhoeffeni Lohmann in combination with a previously published model can be used to give a good approximation of clearance rates. The model is based on tail beat frequency, time spent feeding and subsequent measurements of individual size (trunk length). These estimated clearance rates are in close agreement with clearance rates determined in the laboratory under static and flow-through conditions. A comparison of various techniques (such as measurement of gut pigment, and various particle removal and particle uptake methods) with the behavior-model approach, revealed a convergence of all rates within a threefold range. Most of this discrepancy can be explained by the number of non-feeding individuals and the deterioration of the feeding filters in the field. The main reasons for the similarity of the various clearance rate estimates are the low variability of the behavior of appendicularians in response to environmental variables, such as temperature and particle concentration, and their non-selective retention of particles. The suggested mechanistic approach has great value for estimating the flux of material and energy through populations of appendicularians for which only size and abundance data exist, or for appendicularian species that cannot be assessed empirically. Received: 26 August 1997 / Accepted: 28 September 1998     

Growth,production and ecological significance of Ophiura albida and O. ophiura (Echinodermata: Ophiuroidea) in the German Bight

Growth and production of the shallow-water ophiuroids Ophiura albida and O. ophiura were investigated at two stations in the German Bight from 1988 to 1991. Growth rings visible on the vertebral ossicles of the ophiuroid arms were interpreted as annual age markers. A correction for overgrown first rings allows for more exact estimations of growth and age. In both species growth could be described by Von Bertalanffy growth functions with the asymptotic disc diameter D =10.1 mm, K=0.229 and t _o=-0.192 in O. albida and D =27.7 mm, K=0.084 and t _o=0.042 in O. ophiura. Somatic production was calculated from mass specific growth rates. Annual production:biomass (P:B) ratios were estimated at 0.32 for O. albida and 0.43 for O. ophiura.AWI Publication Number: 618     

Pattern of planktonic bioluminescence in the northern Sargasso Sea: Seasonal and vertical distribution

The vertical structure of bioluminescence potential (BPOT) and flash density (FD) were measured on five cruises to the northern Sargasso Sea in 1987 and 1988. Depth-integrated (0 to 150 m) BPOT did not vary seasonally, remaining within the range 9 to 15 × 10¹⁵ photons m^–2 in all months sampled. Conversely, depth-integrated FD was significantly higher (> 2 × 10⁵ flashes m^–2) during winter (November and March) than during summer (< 9 × 10⁴ flashes m^–2 in May and August). The vertical patterns of BPOT and FD were well correlated within a single profile, more highly so in summer than in winter. Despite intracruise variability in the vertical pattern of BPOT and FD, there were clear summer-winter differences in the vertical distribution of BPOT and FD. During winter, BPOT and FD were maximal and relatively uniform throughout the surface mixed layer; for example in November they declined sharply within the thermocline at 130 to 150 m. During summer, BPOT and FD were greatest (12 to 25 × 10¹³ photons m^–3 and 600 to 1 200 flashes m^–3, respectively) at subsurface depths. Commonly in summer, the upper depth limit of high BPOT and FD occurred at the base of the surface mixed layer (10 to 40 m) and the lower depth limit was located at the base of the subsurface fluorescence maximum (usually at 100 to 120 m).     

Localization of bioluminescence in the siphonophore Nanomia cara

The base of the tentacle of the developing physonect larva (Nanomia cara) has a bioluminescent region. The ability to produce light in the larva is transitory; this ability first appears at about two days of development and is disappearing by eight days, as the larva begins to feed. Subsequently paired bilaterally symmetrical bioluminescent organs are found on the nectophores and the bracts of the adult colony. In both the larva and the adult, bioluminescence is mediated by a calcium specific photoprotein. In all cases the photocytes lack a green fluorescent protein.     

Red bioluminescence in fishes: on the suborbital photophores of Malacosteus,Pachystomias and Aristostomias

Many deep-sea animals produce blue bioluminescence, but species of three genera of midwater dragonfishes also produce red light. In addition to numerous small body photophores, species of these genera (Malacosteus, Pachystomias and Aristostomias) have large suborbital photophores that emit red light and postorbital ones that emit blue light. Pachystomias microdon additionally has a red-emitting preorbital photophore. Fluorescence measurements from the intact suborbital photophores, and from their exposed cores, confirm the previous hypothesis that the red light emitted by Malacosteus is spectrally altered by a superficial shortwave cutoff brown filter. No such filter is present in the other two genera. Studies of the anatomy of the photophores show that the suborbital photophore of Malacosteus is similar in general organisation to other photophores, including the postorbital photophore. The red-emitting photophores of Pachystomias and Aristostomias, however, have a unique organisation, in which the bulk of the photophore comprises a gland that lies outside the thick reflector. Strands of tissue run from the gland into the photogenic core of the photophore through numerous pores in the reflector. The production and use of red light by these fishes is discussed in the context of these results.Electronic Supplementary Material Supplementary material is available for this article at     

Flash kinetics and spatial patterns of bioluminescence in the copepodGaussia princeps

Kinetics and local spatial patterns of bioluminescence in the mesopelagic copepodGaussia princeps (T. Scott), collected by net off the coast of southern California, USA, were investigated from 1986 to 1989. Luminescence elicited by single electrical stimuli began as a localized secretion from any of 16 widely distributed sites on the body. Appendage or body movements swept the particulate luminescent exudate away from the cuticle. Three types of single flashes were identified in response to single electrical stimuli, including a fast flash of ca. 2 s duration, a long flash of 7 s duration, and a 17 s slow flash. In addition, in 20% of the tests a fourth, two-component flash consisting of a fast or long component followed by a slow component was elicited. Frequency of the four flash types was dependent on the recovery time after collection. Shorter, less intense fast and long flashes occurred early in the recovery period, and longer, brighter slow and compound flashes became more common as recovery progressed. The laboratory handling protocol strongly affected total stimulable light (TSL). Strong handling resulted in a reduced TSL, which recovered to the original level upon termination of the stimulation. TSL and flash kinetics in the laboratory were highly dependent on the chronic laboratory stimulation protocol and represent an equilibrium between recovery of luminous reserves and re-stimulation. Flash kinetics and TSL were also affected by the frequency of stimulation. Higher stimulation rates resulted in greater temporal summation of the flashes, a decreased response duration, and a lower TSL. Stimulus frequency and duration therefore determine the nature of the bioluminescent response. Flash kinetics and TSL made with either in situ bathyphotometric measurements of laboratory measurements strongly depend on the extent of prestimulation, depletion of bioluminescent reserves, and the level of stimulation inherent with the particular device.     

Post-larval development of the common north-west European brittle stars Ophiura ophiura,O. albida and Acrocnida brachiata (Echinodermata: Ophiuroidea)

The post-larval skeletal plate ontogeny of the ophiuroids Ophiura ophiura (Linnaeus) and Acrocnida brachiata (Montagu) from the recently metamorphosed 0.3 mm disc diameter size to a juvenile size of 2.0 mm disc diameter has been examined using scanning electron microscopy. Post-larval development of O. albida Forbes is briefly described for sizes between 1.0 and 2.0 mm disc diameter. Early post-larval development in the 0.2 to 1.0 mm discsize range involves considerable morphogenetic change and necessitates the use of a developmental series in the identification of newly settled ophiuroids. Comparison of these ophiuroids with other closely related species reveals that the post-larvae are readily distinguishable at even the smallest sizes by their transient post-larval morphologies. Abbreviated larval development in A. brachiata is suggested from the egg size and post-larval form. The ophiuroid post-larvae were collected from the shallow sublittoral of Oxwich Bay, Bristol Channel, UK, during the years 1980 to 1982.