Asexual reproduction of the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea)

The goal of this research is to enhance our knowledge of thecontributions of doliolids to the planktonic community as consumersand secondary producers. The objectives are to quantify asexualreproduction and carbon release rates of Dolioletta gegenbauriphorozooids at four food concentrations and four temperaturesin order to determine their impact as producers throughout thewater column. Although doliolids are abundant in numerous regionsof the coastal ocean, and are considered to be major planktonicgrazers, data on rates of asexual reproduction are scarce. Laboratoryexperiments were conducted at 20°C at food concentrationsof 7, 20, 60 and 160 µg C l^-1 of Thalassiosira weissflogiiand Rhodomonas sp., and at 60 µg C l^-1 at 16.5, 20, 23.5and 26.5°C, to quantify the phorozooid release rates ofgonozooids, and the amount of carbon released. Results fromthese experiments suggest that release rates increased at 20°Cas concentration increases from 7 to 160 µg C l^-1. Releaserates remained similar as the temperature increased from 16.5to 26.5°C at a phytoplankton concentration of 60 µgC l^-1. Food concentration and temperature has an effect on phorozooidreproductive longevity, size of gonozooids released, and theamount of carbon released asexually. Doliolid reproduction ratesare a function of environmental food concentrations and temperatures,and the results imply that doliolids can be important secondaryproducers in the neritic environment.     

Grazing rates for three life history stages of the doliolid Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea)

Grazing rates for Dolioletta gegenbauri averaged 2.5 ml zooid^–1h^–1 for trophozooids, 2.7 for phorozooids and 5.8 forgonozooids. In situ and laboratory rates were comparable forboth trophozooids and phorozooids.     

Laboratory-measured grazing and ingestion rates of the salp, Thalia democratica Forskal, and the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea)

Grazing and ingestion rates of laboratory-born Thalia democraticaaggregates and Dolioletta gegenbauri gonozooids, phorozooidsand oozooids were determined while fed Isochrysis galbana (4–5µm diameter) alone or in combination with Peridinium trochoideum(16–18 µm diameter) at concentrations of 0.15–0.70mm³ x 1^–1. Grazing rates (ml x zooid^–1 x 24 h ^–1)ranged from 10 to 355, and at zooid weights greater than 5 µgcarbon were in order oozooid > gonozooid > aggregate.Grazing rates increased exponentially with increasing zooidweight. Weight-specific grazing rates (ml x µgC^–1x 24 h^–1) were independent of the four-fold initial foodconcentration. Mean weight-specific grazing rates increasedlinearly with increasing zooid weight for the aggregates andoozooids, but gonozooid mean rates were independent of zooidweight. Aggregate and gonozooid ingestion rates (10⁶ µm³x zooid^–1 x 24 h^–1) ranged from 4 to 134 while oozooidrates ranged from 3 to 67. All ingestion rates were independentof the initial food concentration but increased linearly withincreasing zooid weight at similar rates. All mean weight-specificingestion rates (ml x µgC^–1 x 24 h^–1) wereindependent of zooid weight. The mean aggregate daily ration(µgC ingested x µg body C^–1) was 59% and themean doliolid ration was 132%. Field studies indicate that normalconcentrations of D. gegenbauri in the Georgia Bight clear theirresident water volume (1 m³) in about 4 months, but that highlyconcentrated, swarm populations which occur along thermohalinefronts clear their resident water volume in less than 1 day. ¹Current address: MacLaren Plansearch Ltd., P.O.Box 13250, sta.A.,St.John's, Nfld. A1B 4A5     

Feeding rates of the doliolid, Dolioletta gegenbauri, on diatoms and bacteria

Wild doliolids fed equally efficiently on radiolabelled bacteria>5 µm in size and diatoms >100 µm in size.Feeding by attached asexual buds accounted for 80% of the clearancerates of phorozooids and oozooids     

Lipid composition of the pelagic tunicate Dolioletta gegenbauri (Tunicata, Thaliacea)

The free-swimming sexual stage (gonozoid) of the doliolid, Doliolettagegenbauri, was abundant at a site in the NE Atlantic (58W,20N) during June 1996. Lipid analysis indicated that theseorganisms were comprised mainly of polar lipid with only lowlevels of neutral storage lipid (triacylglycerols) and thisis consistent with a life history strategy of rapid somaticgrowth and gamete production. Fatty acid analysis of the gonozoidssuggested a diverse microplanktonic diet, comprising diatoms,flagellates and dinoflagellates. The fatty acid compositionof D.gegenbauri was dominated by 20:5(n–3) and 22:6(n–3),and is of high nutritional value to predators, particularlyfish. The results are discussed in relation to the organisms'life history strategy and their potential nutritional impacton higher trophic levels.     

Iodine binding and peroxidase activity in the endostyle of Salpa fusiformis,Thalia democratica,Dolioletta gegenbauri and Doliolum nationalis (Tunicata,Thaliacea)

Summary The protothyroid region in the endostyles of four species of tunicates was examined by means of autoradiography and cytochemistry, at both the light and electronmicroscopic levels. To reveal the primary binding site for iodine, autoradiography was carried out on endostylar tissue from animals that had been incubated with high activity ¹²⁵I^- over a short period of time. The specific iodine binding enzyme, a peroxidase, was traced by its reaction with DAB. In accordance with previous findings, the iodinebinding cells proved to be the same as those containing the peroxidase. There were also strong indications of a secondary uptake of iodinated compounds and subsequent release into the body fluid. Together with the ultrastructural features, the data provided strong evidence indicating that these cells constitute a protothyroid region, which partly functions as an endocrine organ, possibly homologous with the vertebrate thyroid gland. Since the number of zones varied between the species, the numeration of the protothyroid region also varied. However, in all the examined endostyles, the protothyroid region was seen to be situated dorsolaterally to the glandular regions of the endostyle concerned with food capture.     

Diet and trophic interactions of a circumglobally significant gelatinous marine zooplankter,Dolioletta gegenbauri (Uljanin, 1884)

Gelatinous zooplankton play a crucial role in marine planktonic food webs. However, primarily due to methodological challenges, the in situ diet of zooplankton remains poorly investigated and little is known about their trophic interactions including feeding behaviour, prey selection and in situ feeding rates. This is particularly true for gelatinous zooplankton including the marine pelagic tunicate, Dolioletta gegenbauri. In this study, we applied an 18S rRNA amplicon metabarcoding approach to identify the diet of captive‐fed and wild‐caught D. gegenbauri on the midcontinental shelf of the South Atlantic Bight, USA. Sequencing‐based approaches were complimented with targeted quantitative real‐time polymerase chain reaction (PCR) analyses. Captive‐fed D. gegenbauri gut content was dominated by pico‐, nano‐ and micro‐plankton including pico‐dinoflagellates (picozoa) and diatoms. These results suggested that diatoms were concentrated by D. gegenbauri relative to their concentration in the water column. Analysis of wild‐caught doliolids by quantitative real‐time PCR utilizing a group‐specific diatom primer set confirmed that diatoms were concentrated by D. gegenbauri, particularly by the gonozooid life stage associated with actively developing blooms. Sequences derived from larger metazoans were frequently observed in wild‐caught animals but not in captive‐fed animals suggesting experimental bias associated with captive feeding. These studies revealed that the diet of D. gegenbauri is considerably more diverse than previously described, that parasites are common in wild populations, and that prey quality, quantity and parasites are likely all important factors in regulating doliolid population dynamics in continental shelf environments.     

On the Pharyngeal Feeding Filter of the Salp Pegea confoederata (Tunicata: Thaliacea)

In Pegea, scanning electron microscopy of what appear to be the least damaged portions of the filter shows that it has a regular rectangular mesh consisting of thick (100 nm) fibres at right angles to thinner (50 nm) fibres. The rectangular pores of the filter are around 3.3 × 0.57 μm. These measured values from filters that have suffered shrinkage (to an uncertain degree) during preparation are considered to indicate that the actual pore size in life is some 4.0 × 0.7 μm. The mucous-net feeding filter of salps differs from that of other tunicates since flow through it results from muscular activity. Calculations based on the estimated pore size and filtering rate suggest that during part of the filtering cycle, the pressure drop across the filter is considerably greater than that across other tunicate mucous-net filters.     

A monograph of the order Pyrosomatida (Tunicata, Thaliacea)

The taxonomy of the order Pyrosomatida is revised based on theexamination of over 300 samples from all parts of the worldoceans. The order consists of a single family (Pyrosomatidae),two subfamilies (Pyrostremmatinae and Pyrosomatinae), threegenera (Pyrostremma, Pyrosomdla and Pyrosoma) and 8 species(Pyrostremma spinosum, P. agassizi, Pyrosomella verticillata.P. operculata, Pyrosoma atlanticum, P. aherniosum, P. ovatumand P. godeauxi n.sp.). The species are briefly described anddata on their distribution are provided. A key to the speciesis given. A cladistic analysis of the distribution of relevantcharacter states is made and ideas on evolutionary developmentare put forward. Biogeographical data are summarized and inconjunction with that the likely modes of speciation in thisgroup are briefly discussed.     

Jet propulsion in salps (Tunicata: Thaliacea)

This paper describes the locomotion of salps by jet propulsion, from a combination of measurements of chamber pressures, static thrust, and electromyographic activity, with kinematic records of free-swimming and tethered salps. From such measurements, estimates are made of the thrust exerted, the drag incurred, and the work performed by single salps, and by chains of linked individuals. It is concluded that salp jet propulsion is a more economical process than is jet propulsion in other animals.     

Distribution and vertical migration of salps (Tunicata, Thaliacea) near Bermuda

Sampling near Bermuda by trawling and SCUBA diving yielded 16species of salps in August 1989 and March-April 1990. Severalspecies occurred in the surface waters only at particular timesof the day. suggesting diel vertical migrations on differentschedules. At the time of sampling, salps were present in lownumerical abundances and constituted only a small fraction ofthe total zooplankton biomass.     

Jet propulsion in Doliolum (Tunicata: Thaliacea)

The jet propulsion of doliolid gonozooids is described from a combination of kinematic, chamber pressure, and muscle electrical records. Doliolids respond to light mechanical stimuli by a rapid contraction of the locomotor muscle bands, producing a single jet pulse which drives the animal forwards or backwards at instantaneous velocities up to 21.4cm·s^?1 (over 50 body lengths·s^?1). Spike potentials from the (multiply innervated) locomotor muscle fibres are variable in size and probably are non-propagating. Maximum chamber pressures during the jet pulse range up to 500 Pa, doliolids ≈4.5 mm long perform around 4 × 10^?6 J work per contraction. Although the locomotor system is specialized for single rapid escape movements, the same movements are used at irregular intervals to maintain the horizontal position of the animal (which is denser than sea water) in the water column. The locomotor system is less economical than that of salps.     

Phosphate excretion rate ofSalpa fusiformis Cuvier (Tunicata : Thaliacea)

The rate of phosphate excretion bySalpa fusiformis was measured for animals of various weights at temperatures of 12.5 to 25 °C. Blastozooids excreted only small, often undetectable quantities of phosphate, in contrast to oozooids, in which the effects of individual weight and temperature were mainly studied. Excretion rate and specific excretion rates were independent of body weight (as total protein). The specific excretion rate increased exponentially with temperature, giving a Q10 of 4.54 (3.17 to 6.43) over the whole temperature range. Using the measurements of Andersen & Nival (1986) on the ammonia excretion rate ofS. fusiformis, we calculated the atomic N/P ratio, which appeared, for oozooids, similar to N/P ratios reported for other zooplanktonic species.     

Structure and ultrastructure of eyes and brains of Thalia democratica (Thaliacea,Tunicata, Chordata)

Salps are marine planktonic chordates that possess an obligatory alternation of reproductive modes in subsequent generations. Within tunicates, salps represent a derived life cycle and are of interest in considerations of the evolutionary origin of complex anatomical structures and life history strategies. In the present study, the eyes and brains of both the sexual, aggregate blastozooid and the asexual, solitary oozooid stage of Thalia democratica (Forskål, 1775 ) were digitally reconstructed in detail based on serial sectioning for light and transmission electron microscopy. The blastozooid stage of T. democratica possesses three pigment cup eyes, situated in the anterior ventral part of the brain. The eyes are arranged in a way that the optical axes of each eye point toward different directions. Each eye is an inverse eye that consists of two different cell types: pigment cells (pigc) and rhabdomeric photoreceptor cells (prcs). The oozooid stage of T. democratica is equipped with a single horseshoe‐shaped eye, positioned in the anterior dorsal part of the brain. The opening of the horseshoe‐shaped eye points anteriorly. Similar to the eyes of the blastozooid, the eye of the oozooid consists of pigment cells and rhabdomeric photoreceptor cells. The rhabdomeric photoreceptor cells possess apical microvilli that form a densely packed presumably photosensitive receptor part adjacent to the concave side of the pigc. We suggest correspondences of the individual eyes in the blastozooid stage to respective parts of the single horseshoe‐shaped eye in the oozooid stage and hypothesize that the differences in visual structures and brain anatomies evolved as a result of the aggregate life style of the blastozooid as opposed to the solitary life style of the oozooid.     

Short communication. Determination of generation time and asexual fecundity of doliolids (Tunicata, Thaliacea)

The goal of this study was to provide temporal information on the generation time of Dolioletta gegenbauri and some of the life cycle's components. At 20C and 90 g C l^-1 of ingestible phytoplankton. D.gegenbauri's life cycle is completed in 20.5 days. Phorozooids 5 mm produce on average 11.0 gonozooids day^-1 over a period of 8-18 days. Utilizing field data on the abundance and size distribution of an assemblage of phorozooids and nurses, in conjunction with experimentally obtained rates, indicates that asexual production per cubic meter by phorozooids with that of nurses should result in rapid colonization of a wide shelf by doliolids, as observed during July and August 1981 on the southeastern continental shelf of the USA.      

Filtration and ingestion rates of Salpa fusiformis Cuvier (Tunicata : Thaliacea): Effects of size,individual weight and algal concentration

Filtration and ingestion rates of Salpa fusiformis Cuvier were determined while fed Phaeodactylumtricomutum Bohlin at concentrations of 2?64 × 10³ cells·ml^?1. Filtration and ingestion rates increase exponentially with increasing length and body protein. The relations between protein content and body length, and between filtration rate and weight are similar for blastozooids and oozooids. A capture efficiency of the order of 6–32% is calculated: Salpa fusiformis seems to have a low retention efficiency, but its very high filtration rate gives it pride of place amongst filter-feeders. Specific filtration rates are independent of weight; specific ingestion rates are independent of weight for blastozooids, but for oozooids they seem to diminish with increasing weight. The mean daily ration (μg C ingested · μg body C^?1) is 107% for a blastozooid and 117% for an oozooid. Specific filtration rates decrease exponentially as particle concentration rises, as for many other filter-feeders, and the specific ingestion rate follows an Ivlev relation.     

Horizontal distribution of Thetys vagina Tilesius (Tunicata, Thaliacea) in the Japan Sea during spring 2004

The occurrence of the salp Thetys vagina was observed in theJapan Sea during spring 2004. Catches up to 187 kg wet weight(WW) per 2.18 x 10⁵ m³ (equal to 0.9 g WW m^–3) were collectedwith 10-m diameter surface-water otter trawl nets. The horizontaldistribution indicated that the high biomass was related tothe area with high chlorophyll a (Chl a) concentration, whichwas located around the subarctic front with the warm TsushimaCurrent. Five prey taxa were identified from the gut contentsof individuals from the high Chl a area. The diatom Coscinodiscusspp. (13–55 µm in diameter) dominated numerically.Another significant prey was the large diatom Coscinodiscuswailesii (219–313 µm) that is an indicator of thespring bloom in this area. The mass occurrence of T. vaginathus appears related to phytoplankton availability, though themechanisms remain uncertain.     

The haemocytes of the salp Thalia democratica (Tunicata,Thaliacea): an ultrastructural and histochemical study in the oozoid

In comparative immunology and evolution of the chordate immune system, tunicates hold an important phylogenetic position as sister group of vertebrates. However, knowledge of the tunicate immune system is limited to the class Ascidiacea, in which some species are now considered model organisms. In the class Thaliacea, represented by fragile pelagic species, the few studies on their haemocytes go back to several decades ago and do not consider comparative aspects with ascidian haemocytes. In this study, we identified various haemocyte types and their distribution in the common salp Thalia democratica by comparative observations under light and electron microscopy and by histochemical, histoenzymatic and immunohistochemical techniques. By comparing specialisations with those of ascidian haemocytes, we detected an undifferentiated cell type (lymphocyte‐like cell) and three categories with four cell types, that is, (i) phagocytic line (hyaline amoebocyte and amoebocyte with large vacuoles), (ii) mast cell‐like line (granular cell) and (iii) storage cells (nephrocyte). Both phagocytes and granular cells appear to migrate in the tunic. Phagocytes adhere to the tunic which internally covers the oral siphon, where they probably function as sentinel cells of the pharynx. Results show the variety of haemolymph cells in the salp similar to phlebobranch ascidians.     

Blooms of tunicates Oikopleura spp. and Dolioletta gegenbauri in the Seto Inland Sea, Japan, during summer

The ecological importance of appendicularians Oikopleura dioica and O. longicauda, and a doliolid Dolioletta gegenbauri as metazoan secondary producers was assessed in summer 1997 in the Seto Inland Sea, Japan. Blooms by the appendicularians occurred twice within a month following peaks of pico-/nanoplankton biomass. The biomass of Oikopleura spp. averaged over the water column (<B> ranged 0.1 to 8.0 μgC l-¹ and temporal average of <B> over the survey period ( ) was 3.2 μgC l-¹, 70% of for calanoid copepods. Furthermore, the temporal average of the production for Oikopleura spp. was 2.4 times higher than that for calanoids, reflecting the extraordinary high growth rates of the appendicularians. D. gegenbauri developed its population following a diatom bloom and <B> reached to 37 μgC l-¹ at the peak time. Although individuals of D. gegenbauri were seldom observed during the first half of the survey, for the doliolid (8.4 μgC l-¹) nearly doubled that for calanoids. These results indicate that the gelatinous tunicates Oikopleura spp. and D. gegenbauri play important roles as metazoan secondary producers in the Seto Inland Sea during summer. This revised version was published online in September 2006 with corrections to the Cover Date.