Biofilm diatom community structure: Influence of temporal and substratum variability

Abstract

Diatoms, which are early autotrophic colonisers, are an important constituent of the biofouling community in the marine environment. The effects of substratum and temporal variations on the fouling diatom community structure in a monsoon-influenced tropical estuary were studied. Fibreglass and glass coupons were exposed every month for a period of 4 days and the diatom population sampled at 24 h intervals, over a period of 14 months. The planktonic diatom community structure differed from the biofilm community. Pennate diatoms dominated the biofilms whilst centric diatoms were dominant in the water column. Among the biofilm diatoms, species belonging to the genera Navicula, Amphora, Nitzschia, Pleurosigma and Thalassionema were dominant. On certain occasions, the influence of planktonic blooms was also seen on the biofilm community. A comparative study of biofilms formed on the two substrata revealed significant differences in density and diversity. However species composition was almost constant. In addition to substratum variations, the biofilm diatom community structure also showed significant seasonal variations, which were attributed to physico-chemical and biological changes in both the water and substratum. Temporal variations in the tychopelagic diatoms of the water were also observed to exert an influence on the biofilm diatom community. Variations in diatom communities may determine the functional ecosystem of the benthic environment.     

Activity of Commercial Enzymes on Settlement and Adhesion of Cypris Larvae of the Barnacle Balanus amphitrite,Spores of the Green Alga Ulva linza,and the Diatom Navicula perminuta

Fouling species produce adhesive polymers during the settlement, adhesion and colonization of new surfaces in the marine environment. The present paper tests the hypothesis that enzymes of the appropriate specificity may prevent biofouling by hydrolysing these adhesive polymers. Seventeen commercially available enzyme preparations designed originally for bulk use in a range of end-use applications were tested for their effects on the settlement and/or adhesion of three major fouling species, viz. the green alga Ulva linza, the diatom Navicula perminuta and the barnacle Balanus amphitrite. The serine-proteases were found to have the broadest antifouling potential reducing the adhesion strength of spores and sporelings of U. linza, cells of N. perminuta and inhibiting settlement of cypris larvae of B. amphitrite. Mode-of-action studies on the serine-protease, Alcalase, indicated that this enzyme reduced adhesion of U. linza in a concentration-dependent manner, that spores of the species could recover their adhesive strength if the enzyme was removed and that the adhesive of U. linza and juvenile cement of B. amphitrite became progressively less sensitive to hydrolysis as they cured.     

Evaluating diatom community composition in the absence of marked limnological gradients in the high Arctic: a surface sediment calibration set from Cornwallis Island (Nunavut,Canada)

Fossil diatom assemblages preserved within the sedimentary record in Arctic lakes provide the potential to reconstruct past changes in important limnological variables. During the summers of 1992 and 1993, we examined previously unstudied freshwater ecosystems on Cornwallis Island, Arctic Canada, with the specific objectives of (1) documenting the limnology and modern diatom assemblages from this region, and (2) determining which environmental variables most influence diatom species distributions. The Cornwallis Island study sites displayed the least amount of variance in measured water chemistry variables in comparison to nearly all of our labs’ previous freshwater surveys in the Arctic. The small limnological gradients precluded the development of a statistically robust diatom inference model, but perhaps more importantly, allowed us to explore variations in diatom composition in the absence of marked variations in water chemistry. Diatom species turnover was minimal, with the most common diatom taxa being Achnanthidium minutissima, Nitzschia perminuta, N. frustulum, with lesser percent abundances of Chaemaepinnularia soehrensis, Navicula chiarae, Psammothidium marginulata, and A. kryophila. A small number of study sites differed from the majority with respect to water chemistry (e.g., coastal sites with high specific conductivities) and habitat availability (e.g., ephemeral ponds with extensive moss habitats), and these sites had markedly different diatom assemblages. These data reinforce previous observations that water chemistry and other climate-related factors are the primary environmental controls influencing diatom distributions at high latitudes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dynamics of diatom colonization process in some rivers influenced by urban pollution (Hanoi, Vietnam)

Periphytic diatom communities’ colonization patterns were studied at three sampling stations of the Red–Nhue–Tolich hydrosystem presenting different urban pollution levels by using artificial substrates for 6 weeks in dry season 2005. Structural characteristics of periphytic diatoms developed on glass substrates at each sampling site were followed and compared. This experiment showed, through various general criteria (total diatom density, dry weight biomass) and specific criteria (relative diatom abundances, indices) that the structure of benthic diatoms developed on these substrates was strongly affected by pollution as early as the second week of colonization. Communities exposed to heavily and moderately polluted sites contained species which are known to be saprophilous or tolerant to organic pollution such as Nitzschia umbonata, Nitzschia palea, Cyclotella meneghiniana, Eolimna minima. Growth inhibition of diatom communities at the heavily polluted site was mostly related to a strong increase of organic load rather than to low metallic input, though metallic (Cd and Zn) burdens have been successfully quantified in the biofilms developed at the three studied sites. Nevertheless, no significant difference in species richness and diversity index between colonization duration times was observed. Based on values of diatom indices Indice de Polluosensibilité (IPS) and diatom assemblage index to organic water pollution (DAIPo), water quality could be classified as medium quality at Red site, polluted at NT₂ and heavily polluted at TL. Thus, the use of diatoms as a tool for water assessment appears suitable for monitoring rivers in Vietnam, as it is in several other countries.     

LIGHT‐INDUCED MOTILE RESPONSES OF THE ESTUARINE BENTHIC DIATOMS NAVICULA PERMINUTA AND CYLINDROTHECA CLOSTERIUM (BACILLARIOPHYCEAE)1

Motility of estuarine epipelic (mud‐inhabiting) diatoms is an important adaptation to living in biofilms present within fine sediments. Motility allows cells to migrate within the photic zone in response to a wide range of environmental stimuli. The motile responses of two species of benthic diatoms to photon fluence rates and spectral quality were investigated. Cultures of Navicula perminuta (Grunow) in van Heurck and Cylindrotheca closterium (Ehrenb.) J. C. Lewin et Reimann both exhibited photoaccumulation at ～200 μmol · m^?2 · s^?1 and photodispersal from photon flux densities (PFDs) of ～15 μmol · m^?2 · s^?1. Photokinesis (changing cell speed) contributed toward photodispersal for both species, and red light (λ = 681–691 nm) was most effective at inducing this process. N. perminuta showed a phototactic (directional) response, with active movement in response to a light gradient. Although this response was exhibited in white light, these directional responses were only elicited by wavelengths from 430 to 510 nm. In contrast, C. closterium did not exhibit phototaxis under any light conditions used in this study. Motile benthic diatoms thus exhibit complex and sophisticated responses to light quantity and quality, involving combinations of photokinesis and phototaxis, which can contribute toward explaining the patterns of large‐scale cell movements observed in natural estuarine biofilms.     

Temperature- and Light-Dependent Performance of the Cyanobacterium <Emphasis Type="Italic">Leptolyngbya Foveolarum</Emphasis> and the Diatom <Emphasis Type="Italic">Nitzschia Perminuta</Emphasis> in Mixed Biofilms

This paper investigates the role of species interactions as a mechanism determining the changing seasonal abundance of microphytobenthic species. Different kinds of interactions can occur between microphytobenthic species, e.g., interference competition (a species directly hindering the growth of another) or resource competition. If such interactions are strong, the capacity of species to exploit parts of the seasonal spectrum of temperature and light conditions could be greatly affected. A model system of two freshwater benthic phototrophs, the cyanobacterium Leptolyngbya foveolarum (Rabenhorst ex Gomont) Anagnostidis et Komárek and the diatom Nitzschia perminuta (Grunow) M. Pergallo, was used to study the capacity of each species to grow in ranges of temperature (7, 15 and 25 °C) and light (5, 40 and 200 μmol m⁻² s⁻¹) conditions in single-species and two-species cultures. Growth was followed for 14–17 days by measuring chlorophyll a and maximum photosynthetic capacity using spectrophotometry and pulse amplitude modulated (PAM) fluorimetry. A PHYTO-PAM fluorimeter facilitated simultaneous observations in mixed cultures on the two species. In mixed cultures, the diatom appeared to be a ‘cool season species’ (low temperature and low light intensity) and the cyanobacterium a ‘summer or autumn species’ (higher temperature and light intensities). This is different than predicted by monoculture experiments, where a wide range of optimal growth conditions was found. Two-species biofilm tests indicated inhibitory effects of the cyanobacterium on the diatom species, especially under conditions favorable to the cyanobacterium. High or low light intensities, increase of local pH caused by depletion of inorganic carbon, and limitation of other inorganic nutrients (resource competition) were examined as factors contributing to diatom inhibition, but none provided an acceptable explanation for observed growth patterns. Our results pointed towards interference competition.     

Epiphytic diatom communities on intertidal seaweeds from Iceland

Epiphytic diatom communities on macroalgae from Iceland coastal waters were investigated during July 2005. Ten species of seaweeds have been collected belonging to brown, red and green algae. The analysis of epiphytic diatom community was carried out under scanning electron microscopy. The epiphytic diatom abundances varied from 7 ± 5 to 7524 ± 3491 cells mm⁻². Erect growth forms were the most abundant, representing on average 50% of the total diatoms (Achnanthes cf. brevipes var. parvula, Tabularia investiens, T. fasciculata, Hyalosira cf. delicatula, Gomphoseptatum aestuarii, Pseudogomphonema plinskii), followed by adnate (29%) (Cocconeis stauroneiformis, C. scutellum) and motile forms (21%) (Nitzschia cf. amphibia and Navicula perminuta). Highly branched seaweeds with articulated thallus surface, offering a number of microenvironments to be occupied by the epiphytes, showed a high level of colonization, mainly due to erect and motile diatoms. Flat thalli with smooth surface allowed for the growth of mainly erect diatoms.     

Nutrient limitation in boreal rich‐fen vegetation: A fertilization experiment

Abstract. Rich‐fen vegetation influenced by hay‐making in the Sølendet Nature Reserve, Central Norway, was fertilized with N, P and K in a full‐factorial fertilization experiment to investigate the nutrient limitation of plant growth at both community and species levels. Above‐ground biomass, shoot density and nutrient concentration were measured in several species and groups of species at three sites after two years of fertilization. At the community level, the results indicate multiple limitation by N and P in the two least productive rich‐fen communities: one characterized by small sedges and herbs, and the other by high abundance of Menyanthes trifoliata and tall sedges. Increased nutrient availability had no effect on a more highly productive, tall‐growing, spring‐influenced community, indicating no nutrient limitation. The results at the species level correspond well with those at the community level, indicating multiple limitation by N and P in most of the dominant and sub‐dominant species. However, P seems to limit growth more than N in Succisa pratensis, and N seems to limit growth more than P in Carex panicea. Furthermore, Eriophorum angustifolium seems to be limited by K. The results did not show which nutrient limits the growth of Carex dioica, C. lasiocarpa and Trichophorum cespitosum. Indications that growth in low‐productive, boreal rich‐fen communities is generally limited by P was not confirmed.     

Diatoms in Acid Mine Drainage and Their Role in the Formation of Iron-Rich Stromatolites

Adverse conditions in the acid mine drainage (AMD) system at the Green Valley mine, Indiana, limit diatom diversity to one species, Nitzschia tubicola. It is present in three distinct microbial consortia: Euglena mutabilis-dominated biofilm, diatomdominated biofilm, and diatom-exclusive biofilm. E. mutabilis dominates the most extensive biofilm, with lesser numbers of N. tubicola, other eukaryotes, and bacteria. Diatom-dominated biofilm occurs as isolated patches containing N. tubicola with minor fungal hyphae, filamentous algae, E. mutabilis, and bacteria. Diatom-exclusive biofilm is rare, composed entirely of N. tubicola.

Diatom distribution is influenced by seasonal and intraseasonal changes in water temperature and chemistry. Diatoms are absent in winter due to cool water temperatures. In summer, isolated patchy communities are present due to warmer water temperatures. In 2001, the diatom community expanded its distribution following a major rainfall that temporarily diluted the effluent, creating hospitable conditions for diatom growth. After several weeks when effluent returned to preexisting conditions, the diatom biofilm retreated to isolated patches, and E. mutabilis biofilm flourished.

Iron-rich stromatolites underlie the biofilms and consist of distinct laminae, recording spatial and temporal oscillations in physicochemical conditions and microbial activity. The stromatolites are composed of thin, wavy laminae with partially decayed E. mutabilis biofilm, representing microbial activity and iron precipitation under normal AMD conditions. Alternating with the wavy layers are thicker, porous, spongelike laminae composed of iron precipitated on and incorporated into radiating colonies of diatoms. These layers indicate episodic changes in water chemistry, allowing diatoms to temporarily dominate the system.     

DIATOMS (BACILLARIOPHYTA) OF ISOLATED ISLANDS: NEW TAXA IN THE GENUS NAVICULA SENSU STRICTO FROM THE GALáPAGOS ISLANDS1

The diatoms (Bacillariophyta) from a coastal lagoon from the Diablas wetlands (Isla Isabela, the Galápagos Islands) were studied in material from surface samples and a sediment core spanning the past 2,700 years in order to examine evidence of diatom evolution under geographic isolation. The total number of taxa found was ～100. Ultrastructural variation in valve morphology between members of Galápagos taxa was used to describe 10 species from the genus Navicula sensu stricto, which are new to science. Four taxa: N. isabelensis, N. isabelensoides, N. isabelensiformis, and N. isabelensiminor, shared several key characteristics that may be indicative of a common evolutionary heritage; these species therefore provide possible evidence for the in situ evolution of diatoms in the Galápagos coastal lagoons. Shared morphological characteristics include: (i) stria patterning in the central area, (ii) an elevated and thickened external raphe‐sternum, (iii) external central raphe endings that are slightly deflected toward the valve primary side, and (iv) an arched valve surface. To explain these findings, two models were proposed. The first suggested limited lateral diatomaceous transport of Navicula species between the Galápagos and continental South America. Alternatively, these new species may be ecological specialists arising from the unique environmental conditions of the Galápagos coastal lagoons, which restrict the colonization of common diatom taxa and enable the establishment of novel, rare species. The Diablas wetlands are an important site for diatom research, where local‐scale environmental changes have combined with global‐scale biogeographic processes resulting in unique diatom assemblages.     

Spatial and temporal variations in moss-inhabiting summer diatom communities from Potter Peninsula (King George Island, Antarctica)

This work represents the first analysis of the spatial and temporal variations in the diversity and community structure of diatoms associated with mosses, either dry or wet, sampled during the summer season from the littoral zone of three ponds in Potter Peninsula (King George Island, South Shetland Islands). It also seeks to explore the effect moisture content in the mosses can have on the structure of the diatom assemblages. The ponds approximately had the same number of species, and differences were found between the dry and wet sampling points. The same 4 species were predominant in the three ponds (Nitzschia alpina, N. perminuta, Staurosira oldenburgiana, and Pinnularia subantarctica var. elongata), but not in the same proportions. The first three species differed from those which were found to be dominant in other basins covered by mosses in southern South America and in the sub-Antarctic region. The latter, P. subantarctica var. elongata, had only been found to be abundant in a moss sample from another South Shetland Island. The diatom assemblages were found to be more strongly related to habitat traits (e.g. distance to the sea, ponds’ areas, and the vegetation’s moisture levels) than to the physical–chemical variables measured in the water.     

Autoinducer‐2 is produced in saliva‐fed flow conditions relevant to natural oral biofilms

Aims: We evaluated the ability of a dual‐species community of oral bacteria to produce the universal signalling molecule, autoinducer‐2 (AI‐2), in saliva‐fed biofilms. Methods and Results: Streptococcus oralis 34, S. oralis 34 luxS mutant and Actinomyces naeslundii T14V were grown as single‐ and dual‐species biofilms within sorbarods fed with 25% human saliva. AI‐2 concentration in biofilm effluents was determined by the Vibrio harveyi BB170 bioluminescence assay. After homogenizing the sorbarods to release biofilm cells, cell numbers were determined by fluorometric analysis of fluorescent antibody‐labelled cells. After 48 h, dual‐species biofilm communities of interdigitated S. oralis 34 and A. naeslundii T14V contained 3·2 × 10⁹ cells: fivefold more than single‐species biofilms. However, these 48‐h dual‐species biofilms exhibited the lowest concentration ratio of AI‐2 to cell density. Conclusions: Oral bacteria produce AI‐2 in saliva‐fed biofilms. The decrease of more than 10‐fold in concentration ratio seen between 1 and 48 h in S. oralis 34–A. naeslundii T14V biofilms suggests that peak production of AI‐2 occurs early and is followed by a very low steady‐state level. Significance and Impact of the Study: High oral bacterial biofilm densities may be achieved by inter‐species AI‐2 signalling. We propose that low concentrations of AI‐2 contribute to the establishment of oral commensal biofilm communities.     

The development of marine biofilms on two commercial non-biocidal coatings: a comparison between silicone and fluoropolymer technologies

The antimicrobial performance of two fouling-release coating systems, Intersleek 700® (IS700; silicone technology), Intersleek 900® (IS900; fluoropolymer technology) and a tie coat (TC, control surface) was investigated in a short term (10 days) field experiment conducted at a depth of ca 0.5 m in the Marina Bandar Rawdha (Muscat, Oman). Microfouling on coated glass slides was analyzed using epifluorescence microscopy and adenosine-5′-triphosphate (ATP) luminometry. All the coatings developed biofilms composed of heterotrophic bacteria, cyanobacteria, seven species of diatoms (2 species of Navicula, Cylindrotheca sp., Nitzschia sp., Amphora sp., Diploneis sp., and Bacillaria sp.) and algal spores (Ulva sp.). IS900 had significantly thinner biofilms with fewer diatom species, no algal spores and the least number of bacteria in comparison with IS700 and the TC. The ATP readings did not correspond to the numbers of bacteria and diatoms in the biofilms. The density of diatoms was negatively correlated with the density of the bacteria in biofilms on the IS900 coating, and, conversely, diatom density was positively correlated in biofilms on the TC. The higher antifouling efficacy of IS900 over IS700 may lead to lower roughness and thus lower fuel consumption for those vessels that utilise the IS900 fouling-release coating.     

Nonadditive effects of consumption in an intertidal macroinvertebrate community are independent of food availability but driven by complementarity effects

Suboptimal environmental conditions are ubiquitous in nature and commonly drive the outcome of biological interactions in community processes. Despite the importance of biological interactions for community processes, knowledge on how species interactions are affected by a limiting resource, for example, low food availability, remains limited. Here, we tested whether variation in food supply causes nonadditive consumption patterns, using the macroinvertebrate community of intertidal sandy beaches as a model system. We quantified isotopically labeled diatom consumption by three macroinvertebrate species (Bathyporeia pilosa, Haustorius arenarius, and Scolelepis squamata) kept in mesocosms in either monoculture or a three‐species community at a range of diatom densities. Our results show that B. pilosa was the most successful competitor in terms of consumption at both high and low diatom density, while H. arenarius and especially S. squamata consumed less in a community than in their respective monocultures. Nonadditive effects on consumption in this macroinvertebrate community were present and larger than mere additive effects, and similar across diatom densities. The underlying species interactions, however, did change with diatom density. Complementarity effects related to niche‐partitioning were the main driver of the net diversity effect on consumption, with a slightly increasing contribution of selection effects related to competition with decreasing diatom density. For the first time, we showed that nonadditive effects of consumption are independent of food availability in a macroinvertebrate community. This suggests that, in communities with functionally different, and thus complementary, species, nonadditive effects can arise even when food availability is low. Hence, at a range of environmental conditions, species interactions hold important potential to alter ecosystem functioning.     

Impact of Urban Pollution from the Hanoi Area on Benthic Diatom Communities Collected from the Red, Nhue and Tolich Rivers (Vietnam)

The effects of urban pollution from Hanoi city on the benthic diatom communities of the Nhue–Tolich river system were studied during the 2003 dry season. Benthic diatoms were allowed to grow on glass slides suspended in the water flow for 4 weeks. To reveal the relationship between water quality and diatom communities, Canonical correspondence analysis (CCA) was used on data concerning relative abundances of diatom species and environmental variables. Two diatom indices, IPS and DAIpo, were applied to evaluate water quality in the three rivers. A total of 291 diatom taxa were found in the Red, Nhue and Tolich Rivers. These were mainly cosmopolitan taxa, with some tropical, subtropical and endemic taxa. The most abundant taxa at the Red site were Aulacoseira granulata, Achnanthidium minutissimum, Encyonema minutum, Navicula recens and other halophilous taxa such as Nitzschia kurzii, Seminavis strigosa, Entomoneis paludosa, Bacillaria paradoxa. Diatom assemblages at the Tolich site consisted mainly of Nitzschia umbonata, Nitzschia palea and Eolimna minima. Diatom density ranged from 660 to 30,000 cells/cm². Environmental variables and diatom assemblage composition at all sites were significantly correlated. Two diatom indices gave similar results and indicate the Tolich River with the lowest values as a highly polluted site.     

Effects of small‐scale environmental variation on metaphyton condition and community composition

1. Metaphyton mats typically originate as benthic algal biofilms that experience higher solar radiation and temperatures, and reduced access to nutrients, once they reach the water surface, but the impacts of these physicochemical changes on metaphyton condition and community composition have received little attention. 2. Using microprobes positioned at 0, 2, 4 and 6 cm depth, we recorded small‐scale differences in water chemistry within metaphyton mats constrained in floating nets during an 8‐week period. Concurrent weekly samples of filamentous algae and their diatom epiphytes were collected from the top, middle and bottom of the mats and were analysed for changes in ash‐free dry mass (AFDM) and chlorophyll‐a, nutrient (N, P, C, Si) content and taxonomic composition. 3. Light intensity, temperature and dissolved oxygen declined both with increasing depth in the mat and over the study period. The autotrophic index (=AFDM/chlorophyll‐a) was greatest at the top of the mats and increased over time; samples also had higher C/P and C/N ratios than samples deeper within the mat. 4. Pithophora was consistently the dominant algal filament throughout the study (representing 85% of all filaments averaged over time and depth); epiphytic diatom cover on Pithophora (calculated as epiphyte area index) declined over time, particularly at the top of the mat. 5. Densities of the diatom epiphytes Gomphonema, Cocconeis and Fragilaria increased with increasing depth within the mat, whereas Cymbella/Encyonema was more common in surface samples. 6. Our results indicate that metaphyton mats are highly dynamic communities, spatially organised in part by small‐scale environmental variation and subject to changes in taxonomic composition following their arrival at the water surface.     

Short-term fluctuations in benthic diatom numbers on an intertidal sandflat in the Westerschelde estuary (Zeeland,The Netherlands)

During the period March–May 1991, sediment samples were taken every two or three days at one intertidal station in the brackish part of the Westerschelde estuary. Quantitative cell counts were made in order to investigate the short-term temporal changes in diatom numbers and assemblage structure.Throughout the whole sampling period, the diatom assemblage was dominated by epipsammic diatoms. Three species, Achnanthes delicatula, Opephora cf. perminuta and Catenula adhaerens on average accounted for almost 67% of all valves counted. The epipsammic diatom fraction showed no significant changes in absolute numbers; its species composition appeared relatively stable. In contrast, epipelic diatom densities significantly increased towards the end of the study period. Species composition within this fraction was less stable. Multivariate analysis (Principal Components Analysis), in combination with multiple regression, indicated that total sky irradiance (on the second and third day preceding sampling) and percentage organic matter were related to the short-term fluctuations of the epipelic diatom fraction.     

OCCURRENCE OF AN ISOPRENOID C25 DIUNASATURATED ALKENE AND HIGH NEUTRAL LIPID CONTENT IN ANTRACTIC SEA-ICE DIATOM COMMUNITIES1

The lipid and hydrocarbon composition of natural populations of diatom communities collected during the austral spring bloom of 1985 in the sea-ice at McMurdo Sound, Antartica was analyzed by TLC-FID, GC and GC-MS. Sea-ice diatom communities were dominated by Amphiprora sp., Nitzschia stellata Manguin and Berkeleya sp. at Cape Armitage; N. stellata, Amphiprora, Pleurosigma, N. kerguelensis (O'Meara) Hasle and some small centric diatoms adjacent to the Erebus Ice Tongue; and Porosira pseudodenticulata (Hustedt) Jouse at Wohlschlag Bay. Lipid distributions of the sea ice diatom communities from the Cape Armitage and Ereus sites were characterized by high concentrations of tracylaglecycerol (triacylglycerolplar lipid = 1.0 to 1.5). The hydrocarbon n-C_21:6, common in temperate diatoms, and an isoprenoid C₂₅ diunsaturated alkene were the dominant hydrocarons detected at these two sites. Hydrogenation of the C₂₅ diene produced the known alkane 2, 6, 10, 14-tetramethyl-7- (3-methylpentyl)-pentadecane. The C₂₅ diene is one of several structurally related hydrocarbons reported in many estuarine, coastal and ocean ic sediments. We propose that certain species of diatoms are a likely source of these alkenes in sediments. The first reported biological occurrence of the C₂₅ diene in the green seaweed Enteromorpha prolifera may have been due to the presence of epiphytic microalgae in the field sample analysed.     

Low‐abundant species facilitates specific spatial organization that promotes multispecies biofilm formation

Microorganisms frequently co‐exist in matrix‐embedded multispecies biofilms. Within biofilms, interspecies interactions influence the spatial organization of member species, which likely play an important role in shaping the development, structure and function of these communities. Here, a reproducible four‐species biofilm, composed of Stenotrophomonas rhizophila, Xanthomonas retroflexus, Microbacterium oxydans and Paenibacillus amylolyticus, was established to study the importance of individual species spatial organization during multispecies biofilm development. We found that the growth of species that are poor biofilm formers, M. oxydans and P. amylolyticus, were highly enhanced when residing in the four‐species biofilm. Interestingly, the presence of the low‐abundant M. oxydans (0.5% of biomass volume) was observed to trigger changes in the composition of the four‐species community. The other three species were crucially needed for the successful inclusion of M. oxydans in the four‐species biofilm, where X. retroflexus was consistently positioned in the top layer of the mature four‐species biofilm. These findings suggest that low abundance key species can significantly impact the spatial organization and hereby stabilize the function and composition of complex microbiomes.