Feeding and growth rates 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 feedingand growth rates of Dolioletta gegenbauri gonozooids at fourfood concentrations and four temperatures in order to determinetheir impact as grazers throughout the water column. Althoughdoliolids are abundant in numerous regions of the coastal ocean,and are considered to be major planktonic grazers, data on ratesof feeding and growth are scarce. Laboratory experiments wereconducted at 16.5, 20, 23.5 and 26.5°C to quantify removalof a 50:50 volumetric concentration of Thalassiosira weissflogiiand Rhodomonas sp. at four different food concentrations of20, 60, 160 and 390 µg C l^–1. Results from theseexperiments suggest that clearance rates are similar at concentrationsfrom 20 to 60 µg C l ^–1, and decrease as the foodconcentrations increase to 160 and 390 µg C l ^–1.The ingestion rates increase over a range of phytoplankton concentrationsfrom 20 to 160 µg C l ^–1, then decrease when abnormallyhigh concentrations of 390 µg C l ^–1 are offered.Clearance and ingestion rates increase as temperature increasesfrom 16.5 to 26.5°C. The exponential growth rates rangefrom k = 0.2–0.7, with the lowest rates occurring at thehighest food concentration. Growth rates increase with increasingtemperature from K = 0.1–0.3 at 16.5°C to 0.45–0.7at 26.5°C. In each case, the small- and medium-sized zooidshad higher growth rates than the larger gonozooids. These resultssuggest that doliolid feeding and growth rates are a functionof environmental food concentrations and temperatures, and implythat they can be important consumers in a changing neritic environment.     

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.     

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.     

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.     

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.     

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     

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.     

In situ measurement of filtering rates of the salp, Thalia democratica, on phytoplankton and bacteria

Salps were gently captured in a perspex chamber by a SCUBA diver,who then injected a suspension of ¹⁴C-labeled phytoplanktonand ³H-labeled bacteria into the chamber. After 0.5 h incubationin situ, filtering rates were estimated from incorporation ofthe two isotopes, and expressed as a function of each salp'ssize. Both bacteria and phytoplankton were grazed by Thalia,the latter at higher rates. Weight-specific grazing rates increasedwith increasing size of salp. Estimated hourly rations rangedfrom 1% of bodily C for small Thalia feeding only on phytoplanktonto 8% of bodily C for large Thalia feeding on all particulateorganic C. The method gave repeatable results for Thalia, butwas unsatisfactory for the pteropod, Cavolinia. ¹Present address: Institute of Marine Resources, A-018, Universityof California, San Diego, Scripps Institution of Oceanography,La Jolla, CA 92093, USA.     

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.     

The developing dorsal ganglion of the salp Thalia democratica,and the nature of the ancestral chordate brain

The development of the dorsal ganglion of the salp, Thalia democratica, is described from electron microscope reconstructions up to the stage of central neuropile formation. The central nervous system (CNS) rudiment is initially tubular with an open central canal. Early developmental events include: (i) the formation of a thick dorsal mantle of neuroblasts from which paired dorsal paraxial neuropiles arise; (ii) the differentiation of clusters of primary motor neurons along the ventral margin of the mantle; and (iii) the development from the latter of a series of peripheral nerves. The dorsal paraxial neuropiles ultimately connect to the large central neuropile, which develops later. Direct contact between neuroblasts and muscle appears to be involved in the development of some anterior nerves. The caudal nerves responsible for innervating more distant targets in the posterior part of the body develop without such contacts, which suggests that a different patterning mechanism may be employed in this part of the neuromuscular system. The results are compared with patterns of brain organization in other chordates. Because the salp CNS is symmetrical and generally less reduced than that of ascidian larvae, it is more easily compared with the CNS of amphioxus and vertebrates. The dorsal paraxial centres in the salp resemble the dorsolateral tectal centres in amphioxus in both position and organization; the central neuropile in salps likewise resembles the translumenal system in amphioxus. The neurons themselves are similar in that many of their neurites appear to be derived from the apical surface instead of the basal surface of the cell. Such neurons, with extensively developed apical neurites, may represent a new cell type that evolved in the earliest chordates in conjunction with the formation of translumenal or intralumenal integrative centres. In comparing the salp ganglion with vertebrates, we suggest that the main core of the ganglion is most like the mes-metencephalic region of the vertebrate brain, i.e. the zone occupied by the midbrain, isthmus, and anterior hindbrain. Counterparts of more anterior regions (forebrain) and posterior ones (segmented hindbrain) appear to be absent in salps, but are found in other tunicates, suggesting that evolution has acted quite differently on the main subdivisions of the CNS in different types of tunicates.     

Mechanism of internal fertilization in Pegea socia (Tunicata,Thaliacea), a salp with a solid oviduct

The ovary of the salp Pegea socia (Bosc, 1802) is located at the end of an atrial diverticulum. The ovary consists of a single oocyte encased in a layer of follicle cells and is connected to the atrial epithelium by an oviduct. Transmission electron microscopy shows that the oocyte lacks a vitelline layer, cortical granules, and yolk granules and that the oviduct lacks a continuous lumen. What previous authors thought was a lumen is a line of dense intercellular junctions running down the center of the oviduct. The sperm nucleus in this species, as in other salps, is elongate. The tubular mitochondrion spirals about the sperm nucleus giving it a corkscrew-shape appearance. Sperm reach the ovary when the oocyte is still at the germinal vesicle stage. Many sperm swim up the atrial diverticulum and burrow through the cells of the atrial epithelium, oviduct, and follicular epithelium. Thus oviduct shortening, which occurs when the oocyte is in the meiotic divisions, is evidently unrelated to sperm moving up the oviduct. All previous authors, who argued either that a continuous lumen is necessary for sperm to move up the oviduct or that sperm bypass the oviduct, were incorrect. © 1994 Wiley-Liss, Inc.     

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.     

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.     

Pyrosoma atlanticum (Tunicata, Thaliacea): grazing impact on phytoplankton standing stock and role in organic carbon flux

Pyrosomas are the large group of pelagic tunicates whose trophicrole in pelagic communities has not yet been sufficiently studied.We ran across a local area of high concentration of the mostwidespread and commonest species of pyrosomas, Pyrosoma atlanticum,450 miles off the Congo river mouth. The following was estimated:gut pigment content, defecation rate, organic carbon and pigmentcontent of fecal pellets, and sinking rate. Based on these dataand the measured number of pyrosomas colonies the grazing impacton phytoplankton and the fecal pellet flux were calculated.During the night swarms of 50–65 mm P.atlanticum removed53% of phytoplankton standing stock in the 0–10 m layer;sparsely distributed pyrosomas consumed only 4%. The grazingimpact in the 0–50 m layer was only 12.5 and <1% respectively.The fecal pellet flux resulting from nocturnal feeding of P.atlanticumwhile swarming made up 1.4–1.6 x 10⁶ pellets m^–210 h^–1 or 305–1035 mg C m^–2 10 h^–1 and1.4 x 10⁵ pellets m^–2 10 h^–1 or 87.4 mg C m^–210 h^–1 while non-swarming. Incubation experiments showedthe rapid degradation of fecal pellets at 23°C: the lossof pigment and carbon content was {small tilde}60–70%after 45 h. We believe that given the sinking rate of 70 m day^–1the main part of fecal material does not leave the upper watercolumn and is retained in the trophic web of the epipelagiclayer.     

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.     

Zooplankton feeding ecology: clearance and ingestion rates of the salps Thalia democratica, Cyclosalpa affinis and Salpa cylindrica on naturally occurring particles in the Mid-Atlantic Bight

Aggregate stages of the salps Thalia democratica, Cyclosalpaaffinis and Salpa cylindrica collected by SCUBA diving in theMid-Atlantic Bight were fed with naturally occurring food assemblages.This is one of the few studies where salps have been fed withnatural food assemblages. The estimated clearance rate for allspecies based on disappearance of chlorophyll varied from 82to 444 mL individual^–1 day^–1. Cell counts showedthat T. democratica mostly ingested carbon from autotrophicnanoflagellates and autotrophic dinoflagellates. Ingestion byS. cylindrica was primarily on larger prey, such as dinoflagellates,while C. affinis ingested auto- and heterotrophic nanoflagellates.All main prey types ingested by salps corresponded to thosethat contributed most to biomass at each experimental station.Thus, salps fed on naturally occurring particles primarily inproportion to prey biomass and to their mechanical capacityto be retained and ingested. Feeding by salps on dinoflagellatesand ciliates implies that they may act not only as potentialcompetitors with microzooplankton, but also as consumers ofthem.     

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.