Structural investigations on the mucus from six species of coral

The chemical composition of the mucus from three hard corals (Acropora formosa, Pachyseris speciosa and Fungia fungites) and three soft corals (Sarcophyton sp., Lemnalia sp., and Cespitularia sp.) collected on the Great Barrier Reef (1982–1985) was determined. Significant variation exists in the composition and structure of the six mucus samples, indicating the absence of a common structure for coral mucus. In all cases protein and/or carbohydrate polymers are the major components of the mucus, and lipids are present only in small amounts. The glycose composition varied between species, with fucose (F. fungites and Lemnalia sp.), arabinose (A. formosa), galactose (P. speciosa) and N-acetyl glucosamine (Sarcophyton sp.) being present in high concentrations. With the exception of Sarcophyton sp. and Lemnalia sp., all mucus samples were acidic and contained significant sulphate but no uronic or sialic acids. The amino acid composition of the mucus samples was not unusual, apart from A. formosa, which contained a high percentage of serine and threonine, and F. fungites, which had high levels of glutamic acid present.     

Sloughed mucus and drag-reduction in a school of Atlantic silversides,Menidia menidia

Atlantic silversides, Menidia menidia, were collected by seine-net from the Newport River estuary, North Carolina, USA and maintained in the laboratory. Direct measurement showed that the amount of mucus sloughed off individual silversides was 567 g h^-1 at a swimming speed of 50 cm s^-1. Individual M. menidia were also allowed to swim in a flowtank filled with 0, 1, and 10 ppm of the synthetic drag-reducer Polyox to determine if any changes in tailbeat frequency or amplitude could be correlated with the level of this sloughed material. Tailbeat measurements between seawater controls and either Polyox concentration did not differ significantly. Calculations based on this evidence indicate that the amount of solubilized mucus sloughed off a school of 10 000 M. menidia would be at least two orders of magnitude less than the highest concentration of Polyox tested. We conclude that solubilized mucus in the water column does not confer drag-reduction to schooling M. menidia.     

Effects of diet,ultraviolet exposure,and gender on the ultraviolet absorbance of fish mucus and ocular structures

Ultraviolet (UV) radiation can be damaging to fish skin and ocular components. Coral reef fishes are regularly exposed to potentially harmful radiation. It was recently discovered that tropical marine fishes possess UV-absorbing compounds in their mucus. This experiment demonstrates significant effects of both diet and ultraviolet exposure on the UV-absorbing compounds in the mucus of a tropical wrasse, Thalassoma duperrey. Fish that are exposed to UV radiation increase the UV absorbance of their mucus only if UV-absorbing compounds are provided in their diet. Fish that are protected from UV radiation decrease the UV absorbance of their mucus regardless of diet. Mucus from female T. duperrey absorbed less UV and females had higher rates of skin damage than males. Females sequester UV-absorbing compounds in their pelagic eggs as well as their epithelial mucus, whereas males do not sequester these compounds in the testes. Spectral transmission through the whole eye was not affected by diet or UV manipulations, but corneal tissue transmission decreased significantly in the UV-exposed individuals. These results demonstrate that coral reef fish can adapt to UV exposure, so long as UV-absorbing compounds are available in the diet.Communicated by P.W. Sammarco, Chauvin     

Microheterotrophic utilization of mucus released by the Mediterranean coral Cladocora cespitosa

The amount of mucus released by the Mediterranean coral Cladocora cespitosa (L.) was determined in laboratory experiments and the incorporation of mucus into bacterial biomass was investigated by means of incubation experiments in 1984. Mean mucus release was 8.5 g (mucus dry wt) pclyp^-1 h^-1 and amounted to 44% of the respiratory carbon losses of the coral since mean organic carbon content of freshly collected mucus was 102.2g C mg (mucus dry wt)^-1. Due to the abundance of C. cespitosa in the shallow littoral of the Bight of Piran, the energy content of mucus released is estimated to correspond to about 20% of the phytoplankton primary production in this area. Furthermore, the carbon conversion efficiency of 20% obtained from the bacterial population during decomposition of mucus indicates the high nutritional value of C. cespitosa mucus, although bacterial carbon onto mucus particles contributes less than 0.1% to the total organic carbon pool of the mucus.     

Trail following,speed and fractal dimension of movement in a marine prosobranch,Littorina littorea,during a mating and a non-mating season

We quantitatively studied movement behaviour in the gastropod Littorina littorea in laboratory experiments during periods of their non-mating season (November 1992) and their mating season (April–May 1993). Snails were collected in 1992 and 1993 from one boulder shore on the north west coast of Sweden. In a comparison between the two seasons (one non-mating and one mating) we measured trail complexity of unsexed snails using fractal dimesion, the degree of mucus trail following (coincidence index of marker and tracker trails) and average movement speed of marker and tracker snails. We found no differences in fractal dimension and coincidence index of trails between the two seasons. Tracker snails moved, however, significantly faster than marker snails during both seasons. This could not be explained by trackers, when locomoting, using the mucus trail deposited by the marker to increase their speed, since there was no correlation between coincidence index and tracker speed. During the mating season we also conducted trail complexity, trail following and speed experiments comparing the behaviour of males and females. There was no difference between males and females in the fractal dimension of movement, nor was there any difference between the mean speed of male and female snails, although male marker snails tended to move faster than female marker snails. Males tracking other males, females tracking other females and females tracking males followed trails about equally long distances (i.e., coincidence indices did not differ). In contrast, males following female mucus trails showed a significantly higher degree of trail following than the other sex combinations. This new finding may suggest that females of L. littorea release pheromones in their mucus trails and that males are able to identify them.     

Diurnal lipid and mucus production in the staghorn coral Acropora acuminata

Net ¹⁴C-accumulation into lipids of Acropora acuminata was rapid and increased with light intensity. Dark ¹⁴C-incorporation was less than 1% noon maximum. Structural lipids were the first radioactively labelled lipid types showing linear ¹⁴C-uptake kinetics. Storage lipids showed non-linear, power-curve kinetics for ¹⁴C-uptake. The rate of ¹⁴C-incorporation into triglycerides and wax esters was maximal during early afternoon and at midday, respectively. Electron microscopic evidence is given for zooxanthellae being primary sites for synthesis of lipids which are exuded from chloroplasts and transferred to animal tissues. Free lipid droplets and crystalline inclusions (wax ester) were common in animal tissues, the inclusions being often associated with mucus-producing cells. The diurnal rate of mucus production was constant. However, ¹⁴C-mucus-lipid production showed a light-dependent diurnal pattern and accounted for 60 to 90% total ¹⁴C of mucus during periods of photosynthetically-saturating light. Here, ¹⁴C was primarily associated with wax esters which were always present in the mucus-lipid. ¹⁴C-triglycerides occur in mucus released only during the day. Lipid and mucus synthesis is discussed in relation to the carbon budget of A. acuminata, in which mucus represented a loss of 40% net C fixation.     

Cytochrome c used as a model system to probe the structural properties of benthic mucilage

Benthic mucilage, whether native or artificially fragmented to microscopic dimensions by ultrasonic treatment, was mixed with cytochrome c, which was used as a 12 000 Da polycationic model compound. Cytochrome c binding profiles proved to depend on the aggregation state of the mucilage. The native mucus matrix binds cytochrome c quasi hyperbolically with an apparent affinity constant K = 1 × 10⁶  M ⁻¹. As shown by chemical modification of both the mucilage and cytochrome c, the binding expression is dependent on the availability of both the positive charges on the cytochrome c surface and the negative charges within the mucus matrix. The extent of binding is sensitive to the ionic strength of the medium. The ultrasonic-stabilized mucus fragments display a peculiar binding profile, with an apparent low affinity, abruptly entering into a high affinity binding region. The results suggest that, depending on the mucus to polycation ratio, a polymeric reticulus builds up. This reticulus can accommodate molecules of at least 12 000 Da molecular weight. The results are also discussed with respect to biological implications. Received: 23 April 1997 / Accepted: 24 November 1997     

Mucous sheet formation on poritid corals: An evaluation of coral mucus as a nutrient source on reefs

The production, release, and subsequent consumption of coral mucus on reefs has been portrayed as a potential pathway for the transfer of coral and zooxanthellae production to other reef organisms. However, reported mucus production rates and analyses of nutritional value vary widely. Poritid corals provide a test system to measure mucus production because they produce mucous sheets which can be collected quantitatively. Fluid mucus and mucous sheets were collected fromPorites astreoides, P. furcata, P. divaricata during 1986 and 1987 on reefs in the San Blas Islands, Panama, La Parguera, Puerto Rico and the Florida Keys, USA. Mucus samples were collected from Indo-pacific poritids (P. australiensis, P. lutea, P. lobata, andP. murrayensis) on the Great Barrier Reef during 1985. Biochemical analyses of the fluid mucous secretions, and the derivative mucous sheet, indicate that the mucus is primarily a carbohydrateprotein complex.Porites fluid mucus had a mean caloric content of 4.7 cal mg^–1 ash-free dry weight (AFDW), while mucous sheets contained 3.5 cal mg^–1 AFDW. Sixty-eight percent of the mucous sheet was ash, while fluid mucus was 22% ash. The high ash and low organic contents suggest that mucous sheets have a low nutritional value. C:N ratios varied (range 6.9 to 13.7 for fluid mucus, and 4.8 to 5.9 for mucous sheets), but were similar to typical C:N ratios for marine organisms. Bacterial numbers and chlorophyll a concentrations were higher on mucous sheets than in the surrounding water. Although bacteria aggregate on mucous sheets, bacteria accounted for less than 0.1% of the carbon and nitrogen content of the mucous sheet. Lower C:N ratios in aged mucus, i.e. mucous sheets versus fluid mucus, were attributed to a loss of carbon rather than an increase in nitrogen. Mucous sheet production accounts for a small proportion (< 2% gross photosynthesis) of published values for coral production. In the San Blas Islands, Panama,P. astreoides produced mucous sheets at a rate of 1.5 g C m^–2 y^–1 and 0.3 g N m^–2 y^–1.P. astreoides andP. furcata produced mucous sheets with a lunar periodicity and may provide approximately monthly pulses of carbon and nitrogen to the reef food-web. However, the low annual production rates suggest that mucous sheets make a small contribution to overall energy flow on coral reefs.     

Effects of sedimentation on the gorgonian <Emphasis Type="Italic">Subergorgia suberosa</Emphasis> (Pallas, 1766)

Effects of oceanic and terrestrial sediment suspensions on the behavioral traits activity, colony expansion, and mucus formation of the gorgonian octocoral Subergorgia suberosa (Pallas, 1766) were studied in the laboratory. Increased sediment concentration significantly changed the logits of activity indices (as colony expansion) and was correlated with decreased activity. For both sources of sediments, the activity index decreased as the concentration of sediment increased. The level of decrease was more significance for terrestrial sediments. The activity index showed no clear time pattern for the control group, but a decrease over time occurred in both sediment treatment groups. Activity change was linearly correlated with exposure time and concentration of terrestrial and ocean sediments. The activity indices of polyps were negatively correlated with the formation of mucus sheets (correlation = −0.536, p = 0.048). For both sources of sediments, the frequency of mucus sheet formation (MSF) increased with sediment concentration. Increases in terrestrial sediment loads resulted in more striking effects. The probability for MSF under different sediment groups at different concentrations over time was modeled by regressions. Frequency of polyp activity is suggested as a suitable biomarker to monitor the effects of sedimentation stress.     

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.     

Functional morphology of the podia and ambulacral grooves of the comatulid crinoid Antedon bifida (Echinodermata)

Feeding units — viz. triplets of unequally-sized podia associated with protective lappets — occur all along the pinnules of adult Antedon bifida (Pennant). Small food particles are trapped by direct mucus impingement to the wall of primary and secondary podia (there are no mucus net or mucus thread helping in this process). Large particles are caught by primary and secondary podia which partly curl over them. Small particles accumulate on the collecting podia before being transferred to the groove, while large particles are transferred one by one. Transfer of particles to the groove occurs by wiping the collecting podia on the ciliary tracts against the ciliary current. When active, tertiary podia always paddle against the ciliary current. They serve in bolus formation (mucus embedding of food particles), but do not participate in bolus compaction or propulsion. Elimination of unwanted particles occur through the action of secondary podia whose movements may disrupt the lappet's palisade, thus creating a sideward current that moves particles from the groove to the outside. Typical podial triplets do not occur along brachial and calycinal grooves. Both brachial and calycinal podia function mostly in guiding and regulating particle flow. The feeding structures of early pentacrinoid larvae of Antedon bifida recall to mind those of pinnules of adult individuals. They consist of twenty-five podia arranged in five radial triplets alternating with five interradial pairs. They are similar to the pinnular feeding structures of adults in that they have both collecting (radial) and paddling (interradial) podia.     

An energy budget for Porites porites (Scleractinia)

An energy budget for Porites porites (Pallas) was determined for specimens from 10 m depth on the Fore Reef of Discovery Bay, Jamaica, between July 1984 and July 1985. Evidence for habitual zooplankton ingestion was not obtained, and P. porites appears to be largely autotrophic. Out of the daily photosynthetically fixed energy, 26% is used for animal respiration and growth, 22% for zooxanthellae respiration and growth, and <1% for colony reproduction as mature planulae; 45% remains unaccounted for. Colony respiration, net photosynthesis, colony skeleton and tissue growth, zooplankton ingestion, reproductive effort and energy content of tissues were measured. Energy loss as continuous mucus secretion was not detected, but may occur by an alternative route via mucus tunics, which occur periodically in situ and in the laboratory. The energy budget suggests that a considerable excess of photosynthetically fixed energy is produced on an ideal sunny day at 10 m depth. This surplus may be required for periodic rather than continuous energy demands, or may be essential to survive less-than-ideal days, when net photosynthetic input is reduced.Contribution No. 357 of the Discovery Bay Marine Laboratory, University of the West Indies     

Coral mucus stable isotope composition and labeling: experimental evidence for mucus uptake by epizoic acoelomorph worms

Mucus released by scleractinian corals can act as an important energy and nutrient carrier in coral reef ecosystems, and a distinct isotopic signature would allow following the fate of this material. This study investigates the natural C and N stable isotopic signatures of mucus released by four scleractinian coral genera (Acropora, Fungia, Pocillopora and Stylophora) in comparison with those of suspended particulate organic matter (POM) in seawater of a Northern Red Sea fringing coral reef near Aqaba, Jordan. The natural δ¹³C and δ¹⁵N signatures of coral mucus differed significantly from seawater POM for the majority of seasonal comparisons, but were inappropriate for explicit tracing of mucus in the coral reef food web. Thus, a labeling technique using stable isotope tracers (¹³C and ¹⁵N) was developed that produced δ¹³C values of up to 122 ± 5‰ (mean ± SE) and δ¹⁵N of up to 2,100 ± 151‰ in mucus exuded by Fungia corals. ¹³C and ¹⁵N-enriched compounds were rapidly (within 3 h) and light-dependently transferred from the endosymbiotic zooxanthellae to the mucus-producing coral host. The traceability of ¹⁵N-labeled mucus was examined by evaluating its uptake and potential utilization by epizoic acoelomorph Waminoa worms naturally occurring on a range of scleractinian coral taxa. This tracer experiment resulted in uptake of coral mucus by the coral-associated acoelomorphs and further demonstrated the possibility to trace stable isotope-labeled coral mucus by revealing a new trophic pathway in coral reef ecosystems.     

Ontogenetic feeding shifts in the meiobenthic harpacticoid copepod Nitocra lacustris

¹⁴C-radiolabelling experiments indicate that adult stages of the salt-marsh harpacticoid copepod Nitocra lacustris (Schmankevitsch) receive a large part of their nutrition through the ingestion and assimilation of certain diatoms. An abundance of empty diatom frustules occurs in the gutpellet contents of field-collected individuals. Naupliar stages do not ingest diatoms in the laboratory, and nauplii from the field do not contain frustules in their gut pellets. Ingestion of diatoms in the laboratory first occurs during the second or third copepodite stage. ³H-radiolabel expeiments and grazing experiments using bacterium-sized beads adhering to the diatoms indicated that both adults and nauplii ingest bacteria adhering to the outer mucus coating of the diatoms (and probably ingest the diatom mucus itself). Adults ingest bacteria (and probably mucus exopolymer) coincidently while ingesting diatoms. The nauplii ingest these components by scraping the outer surface of the diatoms. SEM observations indicate that diatoms are not punctured by the nauplii during feeding. While diatom mucus and associated bacteria play an (as yet unquantified) role in the nutrition of the adults, these components may comprise the bulk of food resources for naupliar stages.     

Function of lateral cilia in suspension feeding of lophophorates (Brachiopoda,Phoronida, Ectoprocta)

Suspension feeding by the brachiopod Laqueus californianus (Koch), the phoronid Phoronis vancouverensis Pixell, the ectoprocts Bugula sp., Membranipora villosa Hincks, and Schizoporella unicornis (Johnston), and actinotrocha and cyphonautes larvae was observed. Lophophorates retain particles on the upstream side of the band of lateral cilia, even after particles have moved lateral to the frontal surface. This suggests that these animals utilize an induced local reversal of beat of the lateral cilia for concentration and capture of suspended food particles. Retention of particles can cease while the current past the tentacles continues. Movement of particles down the frontal surface of the tentacles of L. californianus and the ectoproct species confirms previous observations that mucus strands are not used in particle capture. Possible functions of latero-frontal cilia or bristles are considered. Distribution of the feeding mechanism among phyla, clearance rates, and the lack of fusion of tentacles in brachiopods are discussed. The impingement mechanism previously suggested for lophophorates cannot account for the movements of particles observed here.     

A study of mucus from the solitary coral Fungia fungites (Scleractinia: Fungiidae) in relation to photobiological UV adaptation

The ultraviolet (UV)-absorbance spectrum (300 to 360 nm) of mucus obtained from Fungia fungites (L. 1758, collected in Tahiti in 1991) after being exposed to air for up to 5 min was measured, and UV-absorbing compounds were demonstrated to be present in the mucus, with a peak at 332 nm. The concentration of these UV-absorbing compounds was at a maximum in the first 2 min of secretion and decreased thereafter. Concentration was significantly related to the weight of the coral. Also, as corals were adapted to bathymetric levels of UV radiation, mucus concentration of UV-absorbing compounds decreased significantly with increasing depth.     

Dietary study of the midwater polychaete Poeobius meseres in Monterey Bay,California

This study presents the first quantification of the diet of a gelatinous midwater organism on a temporal basis. Using the Monterey Bay Aquarium Research Institute's remotely operated vehicle Ventana, regular collections of the polychaete Poeobius meseres (Heath, 1930) over a 1 yr period (October 1990 to November 1991) in Monterey Bay yielded intact organisms for the study of feeding behavior and quantitative analysis of stomach contents. In situ observations showed P. meseres feeding in two different ways: (1) by deployment of a mucus web in the water column that passively collects particles for consumption; and/or (2) by grasping detrital material in the water column with its ciliated tentacles. Stomach-content analyses showed that P. meseres is primarily coprophagic, its diet being dominated by fecal pellets from euphausiids and copepods. These fecal pellets appear to provide P. meseres with essentially all its carbon. Although fecal pellets were the most important food item volumetrically, P. meseres also consumed large numbers of diatoms and small numbers of dinoflagellates, chrysophytes, radiolarians, foraminiferans and eggs. The diet of P. meseres appears to reflect primary productivity in the surface waters, with different food items predominant in the diet at different times of the year. Pennate diatoms were most abundant in the diet during the fall, centric diatoms were most abundant during the sumnier, and fecal pellets during the winter. The composition of P. meseres diet suggests that this and other midwater gelatinous organisms have a significant role in the remineralization of particles as they sink from the surface to the deep sea.     

Organ-like gonads, complex oocyte formation, and long-term spawning in Periphylla periphylla (Cnidaria, Scyphozoa, Coronatae)

An account is given about the development of the gametes of the holopelagic coronate scyphomedusa Periphylla periphylla (Péron and Lesueur 1809). The gonads of the species are complex and differ from those thus far described in Scyphozoa in having this characteristic composition of trophocytes, follicle cells, gamete-releasing pores, mucus cells, and resorption cells. Our results differ from those of previous coronate studies with respect to the contact of the oocytes with the gonad tissue throughout the duration of development. Among the Medusozoa, follicle cells have thus far been considered as an apomorphy for the Staurozoa, but their presence in the Scyphozoa casts some doubts on this assumption. From morphological structure and examination of the gonads and gametes, it can be deduced that P. periphylla has true organs and that gamete release is continuous. These results supplement our knowledge of the reproductive biology of this mesopelagic species.     

Acute toxicity evaluation of triazophos,deltamethrin and their combination on earthworm,Eudrilus eugeniae and its impact on AChE activity

The acute toxicity of three formula grade pesticides namely, triazophos (an organophosphate, OP), deltamethrin (a pyrethroid) and combined pesticide (triazophos?+?deltamethrin) was determined in earthworm, Eudrilus eugeniae. They were exposed to different concentrations of these pesticides for 48 h by paper contact toxicity method. The LC₅₀ values for triazophos, deltamethrin and of combination were determined as 0.076, 0.031 and 0.065?μg/cm², respectively. To study the sublethal effect of these pesticides, E. eugeniae were exposed to 5% and 10% of LC₅₀ of pesticides for 48 h. Morphological alterations such as coiling, clitellar swelling, mucus release and bleeding followed by body segmentation were observed in exposed earthworm. Acetylcholinesterase (AChE) activity assayed in different regions of body segment exhibited a significant (p?<?0.05) decrease in its activity particularly in the pre-clitellar region as compared to other regions. The altered behavioural responses in pesticides exposed earthworms would have been due to decline in AChE activity of the nervous system.