Microbial communities in northern Adriatic mucilaginous aggregates: insight into the early phase of aggregate formation

Insight into the initial phase of aggregate formation was provided by comparison between bacterial communities from freshly formed aggregates dominated by the epipelic diatom Cylindrotheca closterium and associated water masses. This study was performed from 2000 to 2006 in the northern Adriatic. The chemotaxonomic structures and physiological conditions were inferred from the fatty acid profiles of the cultured bacterial communities of all implicated components, fresh aggregates, their adjacent waters, oligotrophic high-salinity waters and halocline waters. The results showed similarity between bacterial communities of fresh aggregates and oligotrophic high-salinity water, suggesting their common origin and involvement in the formation of aggregates. In contrast, the origin of the water adjacent to aggregates was different from that of the other components but was similar to the halocline layer and was likely derived from northern Adriatic waters. The presence and activity of heterotrophic bacteria belonging to Alteromonadaceae, which are regularly observed on fresh aggregates, suggest that the early phase of aggregate formation corresponds to an abrupt change of environmental conditions due to the mixing of central and northern Adriatic waters. The initial colonisation of fresh mucilage ascribes to Alteromonas an important ecological function in aggregate community development related to the succession of phytoplankton and heterotrophic bacteria.     

Correlation between the presence of Gonyaulax fragilis (Dinophyceae) and the mucilage phenomena of the Emilia-Romagna coast (northern Adriatic Sea)

The formation of massive amounts of suspended mucilaginous organic matter which periodically affects the Adriatic Sea, has been regarded as a complex physico-chemical phenomenon resulting from the production of extracellular material by phytoplankton. Although the exact cause has remained obscure, the mechanism of its formation has usually been considered to be a long-term process, starting after the late winter-early spring blooms, and involving the participation of various algal species, mainly within the diatom group. In this paper we report on the results of a phytoplankton monitoring programme in northern Adriatic seawaters off the Emilia-Romagna coast of Italy which revealed the constant concomitant presence of the dinoflagellate Gonyaulax fragilis (Schütt) Kofoid and mucilaginous formations. In the early stages of the phenomenon the dinoflagellate was clearly observable by microscopic examination in the mucilage, but as the mucilage aged this alga almost completely decomposed and diatom cells increased in number and became predominant. Although characterized by a slow growth rate in culture, in natural seawater G. fragilis was observed to reach cell densities of up to 7.0×10⁶ cells l⁻¹. The results of this study lead us to propose the hypothesis that the appearance of mucilage in the water column of the Adriatic Sea is the consequence of a seasonal growth of this dinoflagellate favoured by specific environmental circumstances.     

The evolution and phytoplankton composition of mucilaginous aggregates in the northern Adriatic Sea

Recurrent occurrences of visible mucilage “clouds” that cover areas up to several hundred kilometres with vertical dimensions of 20–30 m have been recorded in the stratified water column in the northern Adriatic. In the past this was described as “mare sporco” phenomenon. Past studies of the phenomenon indicated that phytoplankton is an important component of mucilage. Our research revealed the composition of phytoplankton assemblages in different types of mucilaginous aggregates collected during the summers of 1997 and 2000 using pigment biomarkers (HPLC). Phytoplankton biomass in the mucilage samples was very high, ranging from 7.9 μg g⁻¹ to 390.8 μg g⁻¹ of chlorophyll a per unit of dry mass of mucilage. The phytoplankton community in the early, loose stage of mucilaginous aggregates was heterogeneous, as indicated by the diversity of detected pigments. The number of phytoplankton groups decreased as the aggregates aged and diatoms increased in relative biomass (up to 92.7%). Phytoplankton biomass in seawater was similar in years with and without mucilage; however, significantly higher contributions to the total biomass of 19′-hexanoyloxyfucoxanthin-containing phytoplankton (prymnesiophytes) were found in the upper 10 m in spring of the “mucilaginous years” (1997 and 2000) followed by prevalence of diatoms in summer. The F_p pigment index used to assess seawater trophic conditions reached lower values in April–May in mucilaginous years (1997 and 2000) compared to non-mucilaginous years (1998 and 1999). We conclude that the role of prymnesiophytes and other small flagellates is crucial for the initial phases of mucilage appearance. Aggregates represent a favourable environment for the secondary development of opportunistic diatoms that foster mucilage formation.     

Influence of maize root mucilage on soil aggregate stability

This study was undertaken to determine the effects of root exudates on soil aggregate stability. Root mucilage was collected from two-month old maize plants (Zea mays L.) Mucilage and glucose solutions were added at a rate of 2.45 g C kg⁻¹ dry soil to silty clay and silt loam soils. Amended soils, placed in serum flasks, were incubated for 42 d with a drying-wetting cycle after 21 d. Evolved CO₂ was measured periodically as well as the water-stable aggregates and soluble sugar and polysaccharide content of the soil. In mucilage-amended soils CO₂ evolution started with a lag phase of 2–3 days, which was not observed in glucose-amended soils. There was then a sharp increase in evolved CO₂ up to day 7. During the second incubation period there were only small differences in evolved C between treatments. Incorporation of mucilage in both soils resulted in a spectacular and immediate increase in soil aggregate stability. Thereafter, the percent of water-stable aggregates quickly decreased parallel to microbial degradation. On completion of the incubation, aggregate stability in the silty clay soil was still significantly higher in the presence of mucilage than in the control. This work supports the assumption that freshly released mucilage is able to stick very rapidly to soil particles and may protect the newly formed aggregates against water destruction. On the silty clay, microbial activity contributes to a stabilization of these established organo-mineral bounds.     

Fatty acid and phytoplankton compositions of different types of mucilaginous aggregates in thenorthern Adriatic

The biomarkers (fatty acid proportions and ratios characteristicof phytoplankton and bacteria) and phytoplankton species insmall (0.5–2.0 cm) and large (0.5–5 m) marine aggregateswere determined in samples collected in the northern AdriaticSea, during a mucilage event in 1997, as well as in 1993, 1994and 1998, when events were not observed. Types of aggregateswere identified according to various biomarker relationships,particularly those related to bacterial and phytoplankton activitiesand changes in the diatom species composition. Aged mucilaginousaggregates (in summer 1997) showed fatty acid proportions (16P/18P,3.9–7.7) characteristic of the highest phytoplankton activities,and also showed the highest bacterial fatty acid proportions(13.3–17.1%) and ratios (C15:br/C15:0, 4.4–6.0).They showed an different diatom community (dominated by Cylindrothecaclosterium) from that in surrounding waters. These characteristicssuggest a continuous renewal of the aggregate organic matter,supporting the hypothesis that aggregate is a ‘selfsustaining’community. In contrast, both freshly formed marine snow, dominantduring periods without mucilage events, and freshly formed mucilaginousaggregates presented biomarker proportions and ratios similarto those of suspended matter (bacterial fatty acids 3–6.7%,C15:br/C15:0 1.2–4.6, 16P/180.4–4.0), and diatomcomposition similar to that of the ambient water. This indicatesthat marine snow sinks more rapidly than large aggregates, beforesignificant changes can occur in its plankton composition. Otheraging aggregate types showed intermediate characteristics, suggestingthat their residence times in the water column were still sufficientto develop organic production–decomposition cycles thatmodified to various extents their biochemical composition.     

Climate Change and the Potential Spreading of Marine Mucilage and Microbial Pathogens in the Mediterranean Sea

Background

Marine snow (small amorphous aggregates with colloidal properties) is present in all oceans of the world. Surface water warming and the consequent increase of water column stability can favour the coalescence of marine snow into marine mucilage, large marine aggregates representing an ephemeral and extreme habitat. Marine mucilage characterize aquatic systems with altered environmental conditions.

Methodology/Principal Findings

We investigated, by means of molecular techniques, viruses and prokaryotes within the mucilage and in surrounding seawater to examine the potential of mucilage to host new microbial diversity and/or spread marine diseases. We found that marine mucilage contained a large and unexpectedly exclusive microbial biodiversity and hosted pathogenic species that were absent in surrounding seawater. We also investigated the relationship between climate change and the frequency of mucilage in the Mediterranean Sea over the last 200 years and found that the number of mucilage outbreaks increased almost exponentially in the last 20 years. The increasing frequency of mucilage outbreaks is closely associated with the temperature anomalies.

Conclusions/Significance

We conclude that the spreading of mucilage in the Mediterranean Sea is linked to climate-driven sea surface warming. The mucilage can act as a controlling factor of microbial diversity across wide oceanic regions and could have the potential to act as a carrier of specific microorganisms, thereby increasing the spread of pathogenic bacteria.     

Mucilage in Cacti: Its Apoplastic Capacitance, Associated Solutes, and Influence on Tissue 5

Mucilage content in the stems of four sympatric cactus speciesvaried from none for Ferocactus acanthodes, 19% by dry weightfor Opuntia basilaris, 26% for Opuntia acanthocarpa, and 35%for Echinocereus engelmannii. Although the mucilage differedchemically among the species (the arabinose content ranged from17% to 51% of the sugar monomers), its relative capacitance(change in relative water content per unit change in water potential)remained about 15 Mpa^–1. The relative capacitance of thewater-storage parenchyma averaged 1·04 Mpa^–1 andwas consistent with the mucilage content, being lowest for F.acanthodes and highest for E. engelmannii. Mucilage isolatedfrom hydrated tissue was accompanied by solutes with an osmoticpressure of about 0·2 MPa. Such associated solutes influencethe water-release characteristics of mucilage and hence itsrole as an apoplastic capacitor. In particular, extracellularsolutes can facilitate the release of appreciable mucilage-boundwater to the cells at tissue water potentials occurring duringthe initial phases of drought. Key words: Echinocereus engelmannii, Ferocactus acanthodes, Opuntia acanthocarpa, Opuntia basilaris, water potential isotherms     

Bathymetric distribution of medusae in the open waters of the middle and south Adriatic Sea during spring 2002

Twenty-eight species of hydromedusae and one scyphomedusa werecollected from the middle and south Adriatic in Spring 2002.Haliscera bigelowi was recorded for the first time in the AdriaticSea and Dicodonium adriaticum for the southern Adriatic. Thecosmopolitan species Rhopalonema velatum, Aglaura hemistomaand Persa incolorata averaged 88% of total medusan abundance.The Mediterranean species Rhabdoon singulare and Solmissus albescenswere numerous at the deeper stations. These species were foundeither in the southern or in middle Adriatic, and their abundanceis greater than in other regions of the Mediterranean Sea. Speciesdiversity and abundance generally were highest in the upper200 m and decreased with depth. Diel vertical migration patternsincluded some species that nocturnally sought shallower or deeperlayers, and some species migrated in both directions. A fewspecies did not show any vertical migration. Although lightlikely triggers vertical movements, data suggest that foragingstrategies also influence the vertical migration of medusae.     

Pelagic organic matter in the Adriatic Sea in relation to winter hydrographic conditions

The spatial distribution of chlorophyll a (Chla), paniculateorganic carbon (POC) and nitrogen (PON), net zoopiankton ash-freedry weight (AFDW), dry weight (DW) and carbon and nitrogen content,in parallel with primary production, was studied during a jointYugoslav cruise in January 1980. The results were interpretedin relation to hydrographic properties, with special emphasison the relationship between pelagic parameters. Spatial distributionof POC, PON and net zoopiankton AFDW and C and N contents showedsimilarities over the major part of the Adriatic, while Chlaand POC showed dissimilarities since higher Chla values wererestricted to the northwestern part of the Adriatic, separatedfrom the rest of the Adriatic by a pronounced thermal frontextending south of the Istrian peninsula. High POC was foundin the centre of the south Adriatic gyre and in the Jabuka Pit,the latter filled with cold water originating from the arealocated south of the thermal front. The areas of high POC werecharacterized by a higher POC recycling time than in other Adriaticregions. High Chla, POC, PON and net zooplankton AFDW and Cand N contents in parallel with increased primary productionwere found in the southwestern part of the Adriatic off Puglia(Italy) affected by a continuous terrigenous supply of nutrients.The observed spatial distribution pattern of pelagic organicmatter suggests that this distribution is mostly influencedby hydrographic conditions and that the assumption of permanentproductivity zones in the Adriatic is not applicable to thewinter situation.     

Copepod succession in two South African estuaries

The seasonal succession of copepod species was studied fromNovember 1976 through October 1978 in the Swartkops and Sundaysriver estuaries. South Africa. Acartia natalensis appeared inthe plankton in spring and reaches maximum abundance duringsummer and autumn. It was replaced by A. longipatella in lateautumn which reached maximum abundance in winter and spring.Cycles of dominance are regulated by the interaction of temperature,salinity and competition between the two species. A natalensis a more tolerant of low salinity and the replacementof A. longipatella by A. natalensis starts in the upper estuaryand spreads seawards. Maximum abundance of A. natalensis isattained in water of lower salinity than in the case of A. longipatella.In the Sundays estuary A. natalensis appeared briefly in thelatter half of the study and in the upper and middle estuaryonly. In the lower estuary A. longipatella was present duringall seasons. This was due to the absence of competition fromA. natalensis, high salinity and lower summer temperatures dueto marine influence. A third species of copepod, Pseudodiaptomushessei functions as a pioneer species, exploiting "new water"after flooding or strong fresh water inflow. High abundancemay therefore occur during any season and no competition betweenP. hessei and either species of Acartia was observed.     

Effects of Salinity on Growth, Water Relations and Ion Accumulation of the Subtropical Perennial Halophyte, Atriplex griffithii var. stocksii

The effects of salinity on growth, water relations, glycinebetainecontent, and ion accumulation in the perennial halophyte Atriplexgriffithii var. stocksii were determined. The following questionswere addressed: (1) What effect does salinity have on growthresponses at different ages? (2) Is A. griffithii an ion accumulator?(3) Does A. griffithii accumulate glycinebetaine in responseto salinity? Atriplex griffithii plants were grown in pots at0, 90, 180 and 360  m M NaCl in sand culture in a plantgrowth chamber and plants were harvested after 30, 60 and 90d. Plant total dry weight was significantly inhibited at 360m M NaCl. Root growth showed a substantial promotion at 90 mM NaCl. The water potential and osmotic potential of shootsbecame more negative with increasing salinity and time of growth.The Na⁺and Cl^-content in both shoots and roots increased withincreases in salinity. Increased treatment levels of NaCl induceddecreases in Ca⁺, K⁺and Mg²⁺in plants. Atriplex griffithii accumulateda large quantity of ions, with the ash content reaching 39%of the dry weight in leaves. Inorganic ion accumulation is significantin osmotic adjustment and facilitates water uptake along a soil-plantgradient. Glycinebetaine concentration was low in roots, andin stems it increased with increases in salinity. Total amountsof glycinebetaine in leaves increased with increases in salinity,and its concentration increased substantially at 360 m M NaCl.Copyright 2000 Annals of Botany Company Atriplex griffithii, glycinebetaine, growth, ions, water relations.     

Mucilage on the Root Surface and Root Hairs of Sorghum: Heterogeneity in Structure, Manner of Production and Site of Accumulation

Small drops of a mucilaginous character near the tip of roothairs were seen by light microscopy in several species of Sorghumand the Sorghum hybrid Vidan. Electron microscopy revealed thatthe drops are formed from at least two distinct substances,both apparently secreted from the endoplasmic reticulum. In addition, a patchy, fibrillar mucilaginous layer, also withat least two components, was found on the cell wall of the roothairs and on the outer wall of ordinary root epidermal cells.Golgi bodies as well as mitochondria take part in its production.As a rule, the mucilaginous patches are colonized by bacteria. Sorghum, root hairs, mucilage     

Mass accumulation of mucilage caused by dinoflagellate polysaccharide exudates in Tasman Bay, New Zealand

At about 20-year intervals, since at least the 1860s, there have been accounts of the accumulation of large quantities of mucilage in the waters off the northern coast of the South Island of New Zealand. On a few occasions these events have been associated with harmful effects, such as the mass mortality of marine fauna and the impediment of fishing activities, though previous attempts to identify the causative organism(s) or the environmental conditions which promote its development have been unsuccessful. In January 2000, there was a moderate re-occurrence of mucilage accumulation in Tasman Bay that led to the identification of the primary origin of this material as the planktonic, thecate, dinoflagellate Gonyaulax hyalina (Ostenfeld et Schmidt). Macro-aggregate accumulation was occurring in the water column when dinoflagellate cell numbers were relatively low. Experimental work on cultures of this organism showed that the dinoflagellate produced transparent exo-polymers (TEPs) composed primarily of galactose and glucose monomers (44.8 and 29.6 mol%, respectively). Analysis of natural and cultured mucilage showed this had a high inorganic content (>60% (w/w)). A substantial proportion (40% (w/w)) of this inorganic fraction was composed of calcium that apparently played an important role in stabilizing the gel. Phosphorus was also enriched in the mucilage. It is hypothesized that the high inorganic N/P ratio observed in the water column at the time of mucilage formation was an effect rather than a cause of this event, which resulted in the scavenging of particulate material from the water column and enrichment of sediments and bottom waters with organic material and phosphorus.     

Seasonal variation in the species composition of tintinnid cilates in Hiroshima Bay, the Seto Inland Sea of Japan

Seasonal changes in the species composition of tintinnid ciliateswere examined based on time-series samples taken at 2 week intervalsover a 3 year period in Hiroshima Bay, the Seto Inland Sea ofJapan. The maximum abundance of total tintinnids over the entireperiod was 5.7 x 10³ indi viduals l^– Among 32 speciesidentified, a consistent seasonal occurrence was recognizedin 22 species. The relationships between various environmentalfactors and the abundance of each species of tintinnids wereanalyzed using principal component analysis From this analysis,the abundance of many tintinnids was revealed to be associatedwith temperature, the <20 µm size fraction of chlorophylla and water column stability, but not with the <20 µmsize fraction of chlorophyll a, nor with salinity. From theseresults, tintinnid species were divided into five associationtypes: species whose abundance increased with increasing temperature,decreasing temperature, nanophytoplankton abundance, increasingwater column mixing, or increasing water stratification coupledwith low temperature.     

Changes in photosynthetic activity of the diatom Phaeodactylum tricornutum in a culture of limited volume

Changes in photosynthetic activity of a marine diatom duringalgal growth were studied with a typical culture medium formarine algae, ASP-2 (l). As the algal specimen, Phaeodactylumtricornutum was used for the experiments. Nitzschia closteriumand Chaetoceros sp. were also supplementarily used. Photosynthetic and p-benzoquinone Hill activities remarkablychanged with time during algalgrowth; with maximum activityfound in cells at log phase. A rapid decrease occurred in theinterphase from the log to stationary phase. The activity changewas not accompanied by variation in photosynthetic pigment content. The low concentration of phosphorus source was suggested asthe main cause for the change. On supply of extra inorganicphosphate, the time length for holding high photosynthetic activitybecame longer; or, the activity of the cells at stationary phaserecovered at least partly even in the dark. Dark recovery wasnot accompanied by either algal growth or an increase in thecontent of photosynthetic pigments. Inactivation of photosynthesis in the stationary growth phaseand activation by added phosphate in the dark were inferredto be due to changes in activities of both the CO₂-fixing andelectron transfer systems. The observed activity change maynot be attributable to a deficiency in inorganic phosphate asthe substrate for photophosphorylation. Similar changes in photosynthetic activity were also observedwith Nitzschia closterium and Chaetoceros sp. (Received January 30, 1971; )     

Cytochemical and Morphological Evidence for the Involvement of the Plasma Membrane and Plastids in Mucilage Secretion in Aloe arborescens

Most of the volume of Aloe arborescens leaves (volumetric density= 0.68±0.08) is occupied by a mucilage tissue. The mucilagesof Aloe species are glucomannans of great medical and pharmaceuticalimportance. An electron microscopical and histochemical studywas carried out, following the development of the tissue andsecretion processes. In contrast to other reports on polysaccharide secretion inplants, no observable activity was found in the Golgi apparatus.In the young stages of leaf development, positive histochemicalstaining for polysaccharides and structural changes were foundmainly in the plastids. In the mature leaves the mucilage cellsremain alive and intact and secretory activity appears to becentered in the plasma membrane. Aloe arborescens, cytochemistry, mucilage, plasma membrane, plastids, secretion     

Studies of the Surfaces of Desert Plant Seeds: II. Ecological Adaptations of the Seeds of Blepharis persica

A study was made of the role of the hydrated mucilage whicharises from the integumentary hairs of Blepharis seeds and inhibitsgermination where there is an excess of water, and of the anatomicaland physiological basis for this phenomenon. It was found thatthis inhibition was not due to bacterial respiration. The appearanceof the dry mucilage in the unimbibed seeds is described fromscanning electron micrographs. The basic helical framework ofthe secondary wall thickenings and the very unusual second-orderframework of trabeculae in the integumentary hair cells aredescribed.     

Mechanism of Pollination in Gladiolus: Roles of the Stigma and Pollen-tube Guide

Gladiolus has a dry type of stigma. Compatible pollen grainsalight and germinate on the receptive surface of the papillae,penetrate the cuticle and grow towards the style through a sub-cuticularpollen-tube guide of mucilage. This is secreted from epidermalcells of the stylodium and style canal. The cuticle, which coversthe pollen tube guide mucilage, is continuous through the stylecanal to the ovary. The wet stigma of Lilium also has cuticulartissue running through the style canal, covering the mucilage.     

The Possible Involvement of Root-cap Mucilage in Gravitropism and Calcium Movement Across Root Tips of Allium cepa L

Roots of Allium cepa L. grown in aerated water elongate rapidly,but are not graviresponsive. These roots (1) possess extensivecolumella tissues comprised of cells containing numerous sedimentedamyloplasts, (2) lack mucilage on their tips, and (3) are characterizedby a weakly polar movement of calcium (Ca) across their tips.Placing roots in humid air correlates positively with the (1)onset of gravicurvature, (2) appearance of mucilage on tipsof the roots, and (3) onset of the ability to transport Ca polarlyto the lower side of the root tip. Gravicurvature of roots previouslysubmerged in aerated water is more rapid when roots are orientedvertically for 1–2 h in humid air prior to being orientedhorizontally. The more rapid gravicurvature of these roots correlatespositively with the accumulation of mucilage at the tips ofroots during the time the roots are oriented vertically. Therefore,the onset of gravicurvature and the ability of roots to transportCa to the lower sides of their tips correlate positively withthe presence of mucilage at their tips. These results suggestthat mucilage may be important for the transport of Ca acrossroot caps. Allium cepa, root gravitropism, root mucilage, calcium, onion     

Chemical Composition of Viscin Mucilage from Three Mistletoe Species--A comparison

The viscin of mistletoes is a unique tissue which serves toprovide strong adherence of the seed of the parasite to thehost branch. The viscin mucilage of three species, Phoradendroncalifornicum and Arceuthobium americanum (Viscaceae) and Phthirusapyrifolia (Loranthaceae), was analysed. The major componentof the mucilage of Pho. californicum and A. americanum was polysaccharidic;of this, neutral sugars comprised the largest portion, but substantialamounts of uronic acids and proteins were also present. Xyloseand arabinose were the most abundant of the neutral sugars,and analysis showed glycine to be the most abundant amino acidas well as the presence of higher-than-average amounts of histidine.In contrast, in the mucilage of Pht. pyrifolia glucose was themost abundant neutral sugar and the protein content was substantiallyhigher. Based on the differences in composition observed amongthe different species the future use of viscin components astaxonomic markers is suggested. Viscin mucilage, mistletoes, Arceuthobium americanum, Phthirusa pyrifolia, Phoradendron californicum, g.l.c., amino acid analysis