Patchy agglomeration as a transition from monospecies to recurrent plankton blooms

We propose a model for explaining both red tides and recurring phytoplankton blooms. Three assumptions are made, namely the presence of toxin producing phytoplankton, the satiation phenomenon in zooplankton's feeding, modelled by a Holling type II response, and phytoplankton aggregation leading to formation of patches. The dynamics of the plankton population is shown to depend on the fraction of the phytoplankton population that aggregates to form colonies and on the number of the latter.     

On conservativity and shattering for an equation of phytoplankton dynamics

A model of phytoplankton dynamics introduced by Arino describes the evolution of aggregates of phytoplankton by a kinetic-type equation composed of terms describing the growth of the aggregates and their splitting, where the latter is modelled by a singular integral operator of the same form as in the classical fragmentation theory. In this paper we shall show that despite the presence of the growth term, the model displays the typical properties of the fragmentation models, in particular, if the fragmentation rate is unbounded as the size of aggregates tends to zero, then there occurs an unaccounted for loss of the phytoplankton though formally nothing is taken out of the system.     

Modeling aggregation and growth processes in an algal population model: analysis and computations

 Aggregation, the formation of large particles through multiple collision of smaller ones is a highly visible phenomena in oceanic waters which can control material flux to the deep sea. Oceanic aggregates more than 1 cm in diameter have been observed and are frequently described to consist of phytoplankton cells as well as other organic matter such as fecel pellets and mucus nets from pteropods. Division of live phytoplankton cells within an aggregate can also increase the size of aggregate (assuming some daughter cells stay in the aggregate) and hence could be a significant factor in speeding up the formation process of larger aggregate. Due to the difficulty of modeling cell division within aggregates, few efforts have been made in this direction. In this paper, we propose a size structured approach that includes growth of aggregate size due to both cell division and aggregation. We first examine some basic mathematical issues associated with the development of a numerical simulation of the resulting algal aggregation model. The numerical algorithm is then used to examine the basic model behavior and present a comparison between aggregate distribution with and without division in aggregates. Results indicate that the inclusion of a growth term in aggregates, due to cell division, results in higher densities of larger aggregates; hence it has the impact to speed clearance of organic matter from the surface layer of the ocean. Received 1 July 1994; received in revised form 23 February 1996     

Analysis of the effects of fragmentation-coagulation in planktology

A theoretical approach is used to investigate the quantitative and qualitative effects of the flocculation and break-up of aggregates of phytoplankton. The importance of these processes in the study of fish recruitment is discussed. Furthermore, results indicate that fragmentation and coagulation dynamics do not play a significant role in the overall evolution of the phytoplankton population.     

Integration of Vibrio vulnificus into Marine Aggregates and Its Subsequent Uptake by Crassostrea virginica Oysters

Marine aggregates are naturally forming conglomerations of larvacean houses, phytoplankton, microbes, and inorganics adhered together by exocellular polymers. In this study, we show in vitro that the bacterial pathogen Vibrio vulnificus can be concentrated into laboratory-generated aggregates from surrounding water. We further show that environmental (E-genotype) strains exhibit significantly more integration into these aggregates than clinical (C-genotype) strains. Experiments where marine aggregates with attached V. vulnificus cells were fed to oysters (Crassostrea virginica) resulted in greater uptake of both C and E types than nonaggregated controls. When C- and E-genotype strains were cocultured in competitive experiments, the aggregated E-genotype strains exhibited significantly greater uptake by oyster than the C-genotype strains.     

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.     

微囊藻群体形成影响因子及机理

富营养化导致的湖泊、水库蓝藻水华带来了一系列环境和生态问题。微囊藻群体形成,在水体表面的聚集是形成水华的重要策略之一。微囊藻群体形成有效抵御了草食性动物摄食、病毒、细菌的侵害,耐受不良环境因子(紫外辐射、高光强、重金属等)能力显著增强。文章探讨了野外微囊藻群体占优势的机制并综述了影响微囊藻群体形成的外源因子及作用机理,包括非生物因子(营养、温度、光照、重金属、微囊藻毒素、乙醛酸)、生物因子(草食性动物、细菌、鱼类、藻类)。基于现有研究,本综述还对未来研究进行展望:(1)非生物因子与生物因子协同效应对藻类形态影响研究; (2)人类活动对浮游藻类形态及动态的影响研究;(3)藻类形态对沉水植物的响应在未来淡水生态系统修复中的应用; (4)促进藻类群体形成的胞外多糖分泌在未来工业生产中的作用。     

Influence of phytoplankton lysis or grazing on bacterial metabolism and trophic relationships

Experimental microcosms were used to study the dynamics of heterotrophic bacterial populations with respect to phytoplankton loss. In a two-stage linked culture system, we artificially separated production and loss processes of a diatom Phaeodactylum tricornutum. In the first (productive) stage, the algae developed axenically and continuously. The outflow was fluxed in two degradation stages, where phytoplankton-derived detritus resulted respectively from: (1) excretion and by-products of phagotrophic organisms (protozoans), and (2) bacterial degradation through bacterial attachment and lysis. According to the phytoplankton decay mode, i.e., lysis or grazing, bacterial adaptations were different. The study of bacterial productivity and aminopeptidase activity showed specific bacterial evolution during the succession of different prey-predator relationships. The occurrence of aggregates allowed nanoflagellates to develop an alternative diet; they fed not only on bacteria, but also on partially degraded phytoplankton detritus, inducing a strong short-cut in the food chain. Sources and controls of extracellular proteolytic activity are discussed. Such experimental approaches are interesting because they separate bacterial lysis and protozoan grazing of phytoplankton, as well as the fates of their corresponding phytoplankton detritus in the microbial food web.     

Temporal dynamics and spatial variation of algae in relation to hydrology and sediment characteristics in the Okefenokee Swamp,Georgia

In a 20-month study, phytoplankton and periphyton chl a, and dry mass of macroscopic algal aggregates in four marshes and a lake within the Okefenokee Swamp (Georgia, USA) were comparable to other wetlands and lake littoral areas. Chlorophyll levels in two marshes were inversely related to water level and phytoplankton at three marshes developed unimodal maxima following macrophyte dieback. Standing stocks in a vernally inundated marsh were greater than a nearby marsh which was permanently inundated; chlorophyll levels displayed longer blooms in the inundated marsh during periods of low rainfall or after drought. Field dynamics, sediment sorption characteristics and algal bioassays suggest that evaporative drawdown stimulates algae by release of nutrients from exposed peat, while high water levels reduce nutrient release from sediments and disperse phytoplankton through flushing. Equilibrium phosphate concentrations of sediments and algal levels were higher at an abandoned rookery than a nearby non-rookery area, indicating nutrient enrichment from residual guano deposits.     

Influence of organic matter quality in the cleavage of polymers by marine bacterial communities

The influence of the quality of organic matter on the hydrolysisof polymers by marine bacteria was investigated in microcosmscontaining aggregates created in rolling tanks. Two types ofmicrocosms were analysed: microcosms type M1 from unalteredseawater and microcosms type M2 from phytoplankton cultures.Kinetics of aminopeptidase, -glucosidase and ß-glucosidasewere measured in the ambient water and the aggregates in thetwo types of microcosms. Bacteria attached to aggregates expressedenzymes with K_m values higher than those of the bacteria inthe ambient water for the three hydrolytic activities analysedin both types of microcosms. Attached bacteria showed higherrates of polymer hydrolysis than free-living bacteria only inmicrocosms M2 created from freshly produced phytoplanktonicmaterial, while free-living bacteria were more active than attachedbacteria in the microcosms M1 containing unaltered seawater.The ratio V_max/K_m, which describes the ability of enzymes tocompete at low substrate concentration, shows that free-livingbacteria are more efficient at dealing with low substrate concentrationsin microcosms derived from natural seawater, where the liquidphase may be depleted of utilizable dissolved organic matter,than in the microcosms derived from phytoplankton cultures.Our data suggest that the hydrolytic activities of both attachedand free-living bacteria are significantly influenced by thequality of the aggregates and the consequences of this influenceare discussed.     

Influence of age of aggregates and prokaryotic abundance on glucose and leucine uptake by heterotrophic marine prokaryotes.

The kinetics of glucose and leucine uptake in attached and free-living prokaryotes in two types of microcosms with different nutrient qualities were compared. Microcosm type M1, derived from unaltered seawater, and microcosm type M2, from phytoplankton cultures, clearly expressed different kinetic parameters (Vmax/cell and K' m). In aggregates with low cell densities (M1 microcosm), the attached prokaryotes benefited from attachment as reflected in the higher potential uptake rates, while in aggregates with high cell densities (M2 microcosm) differences in the potential uptake rates of attached and free-living prokaryotes were not evident. The aging process and the chemical changes in aggregates of M2 microcosms were followed for 15-20 days. The results showed that as the aggregates aged and prokaryotic abundance increased, attached prokaryotes decreased their potential uptake rate and their K' m for substrate. This suggests an adaptive response by attached prokaryotes when aggregates undergo quantitative and qualitative impoverishment.     

Structural transitions and oligomerization along polyalanine fibril formation pathways from computer simulations

The results of a computer simulation study of the aggregation kinetics of a large system of model peptides with particular focus on the formation of intermediates are presented. Discontinuous molecular dynamic simulations were used in combination with our intermediate-resolution protein model, PRIME, to simulate the aggregation of a system of 192 polyalanine (KA(14) K) peptides at a concentration of 5 mM and a reduced temperature of T* = 0.13 starting from a random configuration and ending in the assembly of a fibrillar structure. The population of various structures, including free monomers, beta sheets, amorphous aggregates, hybrid aggregates, and fibrils, and the transitions between the structures were tracked over the course of 30 independent simulations and averaged together. The aggregation pathway for this system starts with the association of free monomers into small amorphous aggregates that then grow to moderate size by incorporating other free monomers or merging with other small amorphous aggregates. These then rearrange into either small beta sheets or hybrid aggregates formed by association between unstructured chains and beta sheets, both of which grow in size by adding free monomer chains or other small aggregates, one at a time. Fibrillar structures are formed initially either by the stacking of beta sheets, rearrangement of hybrid aggregates or association between beta sheets and hybrid aggregates. They grow by the addition of beta sheets, hybrid aggregates, and other small fibrillar structures. The rearrangement of amorphous aggregates into beta sheets is a critical and necessary step in the fibril formation pathway.     

Aggregation of algae released from melting sea ice: implications for seeding and sedimentation

Summary Factors influencing the fate of ice algae released from melting sea ice were studied during a R V Polarstern cruise (EPOS Leg 2) to the northwestern Weddell Sea. The large-scale phytoplankton distribution patterns across the receding ice edge and small-scale profiling of the water column adjacent to melting ice floes indicated marked patchiness on both scales. The contribution of typical ice algae to the phytoplankton was not significant. In experiments simulating the conditions during sea ice melting, ice algae revealed a strong propensity to form aggregates. Differences in the aggregation potential were found for algal assemblages collected from the ice interior and the infiltration layer. Although all algal species collected from the ice were also found in aggregates, the species composition of dispersed and aggregated algae differed significantly. Aggregates were of a characteristic structure consisting of monospecific microaggregates which are likely to have formed in the minute brine pockets and channels within the ice. Sinking rates of aggregates were three orders of magnitude higher than those of dispersed ice algae. These observations, combined with the negligible seeding effect of ice algae found during this study, suggest that ice algae released from the melting sea ice are subject to rapid sedimentation. High grazing pressure at the ice edge of the investigation area is another factor eliminating ice algae released during melting.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Phytoplankton aggregate formation: observations of patterns and mechanisms of cell sticking and the significance of exopolymeric material

Flocculation of ‘sticky’ phytoplankton cells intorapidly sinking aggregates has been invoked as a mechanism explainingmass sedimentation of phytoplankton blooms in the ocean. Phytoplanktonstickiness, defined as the probability of adhesion upon collision,is one key factor determining the potential for aggregate formation.In the laboratory, we examined variation in stickiness in fivespecies of diatoms and two species of flagellates grown in batchcultures. We also investigated the production of paniculatemucus by phytoplankton cells and its role in aggregate formation,and we studied the effects of solute exudates on cell stickiness.Four of the five diatoms investigated were significantly sticky,while one diatom and both of the flagellates were not sticky.Stickiness varied considerably within species. In the diatomSkeletonema costatum, the typical but not entirely consistentpattern was that stickiness decreased with age of the batchcultures. We were otherwise unable to establish consistent relationshipsbetween cell stickiness and the growth stage of the algae, environmentalconcentrations of inorganic nutrients, and abundances of suspendedand epiphytic bacteria. We showed that the diatom S.costatumat times excretes a solute substance that depresses flocculation.This may reduce cell losses from the euphotic zone during thegrowth phase due to flocculation and sedimentation. We demonstratedtwo different mechanisms of phytoplankton aggregate formation.In the diatom S.costatum, the cells are sticky in themselves,and coagulation depends on cell-cell sticking and does not involvemucus. Aggregates are composed solely of cells. Cells of thediatom Chaetoceros affinis, on the other hand, are not in themselvessticky. Transparent exopolymeric particles (TEP), produced bythe diatom, cause the cells to aggregate and coagulation dependson TEP-cell rather than cell-cell sticking. Aggregates are formedof a mixture of mucus and cells. We found several species ofdiatoms and one flagellate species to produce copious amountsof TEP. TEP from some species (e.g. Coscinodiscus sp.) is stickyand may cause other, non-sticky particles to coagulate. Thisemphasizes the potential importance of diatom-derived paniculatemucus for particle flocculation in the ocean.     

Serum proteins enhance aggregate formation of dissociated fetal rat brain cells in an aggregating culture

Summary Dissociated fetal rat brain cells (Day 14.5 of gestation) reaggregated into small cell clusters and formed large aggregates in a medium supplemented with serum or dialyzed serum in an aggregating culture. In contrast, only small aggregates were produced in a serum-free medium. The present results indicated that albumin, fetuin, transferrin, and {ie1031-1}-antitrypsin enhanced the aggregate formation. Small aggregates produced in a serum-free medium elongated neurites when they were cultured within a collagen gel matrix. Total DNA per flask was almost the same in small and large aggregates. Thus, these serum proteins may well play an important role in the adhesion of small cell clusters and cause the formation of large aggregates in this short-term aggregating culture.     

Physiology of purple sulfur bacteria forming macroscopic aggregates in Great Sippewissett Salt Marsh, Massachusetts

Abstract Purple bacterial aggregates found in tidal pools of Great Sippewissett Salt Marsh (Falmouth, Cape Cod, MA) were investigated in order to elucidate the ecological significance of cell aggregation. Purple sulfur bacteria were the dominant microorganisms in the aggregates which also contained diatoms and a high number of small rod-shaped bacteria. Urea in concentrations of ≥ 1 M caused disintegration of the aggregates while proteolytic enzymes, surfactants or chaotropic agents did not exhibit this effect. This suggests that polysaccharides in the embedding slime matrix stabilize the aggregate structure. In addition cell surface hydrophobicity is involved in aggregate formation. The concentration of dissolved oxygen decreased rapidly below the surface of aggregates while sulfide was not detected. The apparent respiration rate in the aggregates was high when the purple sulfur bacteria contained intracellular sulfur globules. In the presence of DCMU, respiration remained light-inhibited. Light inhibition disappeared in the presence of KCN. These results demonstrated that respiration in the aggregates is due mainly to purple sulfur bacteria. The concentration of bacteriochlorophyll (Bchl) a in the aggregates (0.205 mg Bchl a cm⁻³) was much higher than in the pool sediments but comparable to concentrations in microbial mats of adjacent sand flats. Purple aggregates may therefore originate in the microbial mats rather than in the pools themselves. Rapid sedimentation and high respiration rates of Chromatiaceae in the aggregates would prevent the inhibition of Bchl synthesis if aggregates were lifted off the sediment and up into the oxic pool water by tidal currents.     

Shoot meristem differentiation in shoot primordia culture (SPC) obtained from root primordia culture (RPC) of Solanum lycopersicoides Dun.

We present a unique example of conversion of the morphogenesis type (from rhisogenesis to shoot organogenesis) in in vitro cultures of Solanum lycopersicoides. Liquid shoot primordia cultures (SPC) were obtained from root primordia culture (RPC) on two kinds of MS-based media with BA. The first SMS₈, contained a full set of organic compounds; the second, ²SMS₈, was devoid of organic compounds except sucrose and edamine. Two weeks after passage of RPC onto both the media, disintegration of root primordia and cell aggregates began. After 8 weeks of cultivation on SMS₈ and ²SMS₈ media, SPC aggregates developed from meristematic cells located near the vascular tissue of disintegrating RPC aggregates. Initiation of shoot meristems started from meristematic cells centers which were localised under the surface of the newly formed SPC aggregates. The change in the type of morphogenesis occurred faster on medium SMS₈, but the SPC on medium ²SMS₈ was characterized by more frequent formation of shoots and plants (45% of aggregates, in the case of SPC on medium SMS₈, and 90% in the case of SPC on medium ²SMS₈). This fact was correlated with the structural organization of the meristematic centers. Our results indicate that an RPC system has high morphogenetic potential due to the continual presence of meristematic cells. The change in the type of morphogenesis was followed by a rebuilding of the aggregate structure on the basis of the meristematic cells already existing in RPC, which gave rise to SPC aggregates from which shoot meristems developed.     

Dimorphic aggregation behavior of a fusion polypeptide incorporating a stable protein domain (EGFP) with an amyloidogenic sequence (retroCspA)

We describe the behavior of a polypeptide consisting of the genetic fusion of a structurally stable single-domain protein, EGFP (an analog of the green fluorescent protein) with an amyloidogenic sequence, retroCspA (known to readily form amyloid fibrils). Refolding of the fusion protein through single-step removal of denaturant and salt results in precipitation into amyloid aggregates displaying fibrillar morphology, thioflavin T binding as well as green fluorescence. Refolding through step-wise reduction of denaturant concentration in the presence of salt yields a soluble aggregate containing a folded, thermally-stable, non-fluorescent EGFP domain. Together, these results indicate that retroCspA forces the fusion protein to aggregate; however, the EGFP domain remains folded in a native-like structural format in both soluble aggregates and precipitates.     

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.     

马尾松人工林土壤各粒径团聚体湿筛后的有机碳分配

选取25a、45a和65a马尾松人工林为研究对象,采用湿筛法对各粒径土壤团聚体分别湿筛。探究了马尾松人工林各粒径团聚体湿筛后的团聚体有机碳分配,以探讨各粒径团聚体湿筛后分配到同一粒级团聚体有机碳含量及其对团聚体水稳性的贡献差异。结果表明：种植年限增加显著降低土壤团聚体水稳性（P<0.05）;各粒径团聚体湿筛后分配的有机碳随粒级减小含量呈先降后增趋势,以保持原粒级团聚体有机碳（12.96-32.01 g/kg）含量最高,其次是<0.25 mm粒级（8.08-23.53 g/kg）。各粒径团聚体湿筛分配到同一粒级的有机碳以保持原粒级的含量最高（P<0.05）;土壤团聚体水稳性与各粒径团聚体湿筛后保持原粒径的有机碳呈显著或极显著正相关（P<0.05或0.01）,分配到越小的粒级正相关性越不显著。此外,团聚体水稳性与各粒径团聚体湿筛分配到同一粒级的有机碳呈正相关,以保持原粒级相关性最高（P<0.01或0.05）;回归方程及相关性系数表明,有机碳与保持原粒径团聚体呈显著呈或极显著正相关（P<0.05或0.01）,与消散到其他粒级的团聚体呈负相关或极显著负相关（P>0.05或<0.01）。本研究得出有机碳含量增加促进更大粒径团聚体形成。反之,促使大粒径团聚体向较小粒径团聚体转化。同一粒级团聚体间,保持原粒级团聚体比易转化形成更大粒级团聚体有更高的有机碳含量和更强的水稳性,这对团聚体的固碳提供了新的认识。