Symbiotic Relations of Sediment-Agglutinating Nematodes and Bacteria in Detrital Habitats: The Enzyme-Sharing Concept

Abstract. A new concept (‘enzyme sharing’) concerning the interaction of marine nematodes and microbes in the degradation of sedimentary detritus is presented. Elements of this concept are (1) the notorious tendency of many aquatic nematodes to agglutinate detrital particles by mucus secretions, (2) new observations on the stimulation of microbial growth by nematodes in agar plates, and (3) literature data on limited endogenous proteolytic capacities of aquatic nematodes.
Observations on nematode‐microbe associations in agar plates prompted the conceptual synthesis. In agar medium without the addition of any nutrients a spectacular growth of bacteria was visible on the sinusoidal crawling trails of nematodes only 2 – 3 days after introduction of the worms (species of Adoncholaimus, Anoplostoma and Sabatieria). Juveniles of Anoplostoma that hatched in the agar cultures left their minute trails in the medium and these were rapidly occupied by bacteria. The nematodes repeatedly visited their bacterial trails, which persisted as a peculiar biotic structure for more than one year and survived the nematodes.
In sterile agar preparations containing the fluorogenic methylumbelliferyl‐β‐glucoside in the presence of the nematode Adoncholaimus, an enhanced fluorescence of the medium was visible, indicating β‐glucosidase activity. We therefore assume that oncholaimid nematodes discharge enzymes that alone, or in concert with microbial activities, contribute to the hydrolytic cleavage of refractory polysaccharides containing β‐glucosidic bonds such as agar components and cellulose. The sugars thus produced may then be taken up by the nematodes and concomitantly support the conspicuous growth of microbes. Since we did not observe any feeding of the nematodes on the associated microbes in agar plates, we question the nutritive potential of intact microbial cells for a number of nematodes abounding in detrital habitats, and call attention to the significance of ambient dissolved or adsorbed organic monomeric nutrients.
Consequently, we perceive the puzzling perpetual accretion of detrital organic particles to sediment agglutinations by nematodes as an adaptation for operating an ‘enzymatic reactor’ for the production of dissolved nutrients. We hypothesise a relationship of mutual commensalism of nematodes and heterotrophic microbes in detrital habitats and propose the term ‘enzyme sharing’ for this relationship. Both parties invest in a common enzyme pool that decomposes organic detritus for their nutrition. We present here evidence that nematodes contribute β‐glucosidase, which is involved in the cellulase system. Data from the literature suggest that microbial enzymatic processing of detrital proteins yields amino acids available to nematodes, which apparently have no efficient proteolytic enzyme system in their intestines.     

Microbial Biomass and Organic Nutrients in the Deep-Sea Sediments of the Central Indian Ocean Basin

In order to assess the impact of deep-sea mining on the in situ benthic life, we measured the microbial standing stock and concentration of organic nutrients in the deep-sea sediments of the Central Indian Ocean Basin in the Indian pioneer area. Sediments were collected using box core and grab samples during September 1996. The total bacterial numbers ranged from 10 10 -10 11 cells per g -1 dry weight sediment. There was a marginal decrease in the number of bacteria from surface to 30 cm depth, though the subsurface section registered a higher number than did the surface. The highest numbers were encountered at depths of 4-8 cm. The retrievable number of bacteria were two orders less in comparison with the direct total counts of bacteria. An almost homogeneous distribution of bacteria, total organic carbon, living biomass, and lipids throughout the depth of cores indicates active microbial and benthic processes in the deep sea sediments. On the other hand, a uniform distribution of total counts of bacteria, carbohydrates, and total organic carbon in all the cores indicates their stable nature and suggests that they can serve as useful parameters for long-term monitoring of the area after the benthic disturbance. Further studies on temporal variability in this region would not only verify the observed norms of distribution of these variables but would also help to understand restabilization processes after the simulated benthic disturbance.     

Food Preferences of Dominant Salt Marsh Herbivores and Detritivores

Abstract. Food preferences of six dominant salt marsh invertebrates were studied to identify detritivores and to assess differences in their diets. Animals fed on agar suspensions of dead and live foods in petri dishes with four compartments. Only two foods were included in each dish. Relative palatability was assessed by measuring the amount of suspension removed or by counting the number of feeding marks left on the surface of the suspensions.
Feeding marks reflected differences in mouth parts and feeding mechanisms of the six invertebrates. Melampus bidentatus, Orchestia grillus , and Philoscia vittata preferred dead litter over live tissues of marsh graminoids, blue-green algae, and sulfur bacteria. No preference for detritus from different graminoids was shown by M. bidentatus. Orchestia grillus fed preferentially on Spartina patens detritus. Philoscia vittata preferred detritus from S. alterniflora and S. patens. Blue-green algae and sulfur bacteria were preferred over detritus by Littorina saxatalis , but detritus was preferred over live graminoids. Littorina saxatalis fed preferentially on Juncus gerardi detritus. Blue-green algae, sulfur bacteria, and live graminoids were preferred over detritus by L. littorea. Talorchestia longicornis also preferred blue-green algae.
On the basis of their food preferences, Littorina saxatalis, Melampus bidentatus, Orchestia grillus , and Philoscia vittata were classified as detritivores. Feeding on detritus from different plant species could result in a partitioning of this food resource in the detritus-based food chains of the salt marsh ecosystem.     

Observations of resuspended diatoms in the turbid tidal edge

Observations of resuspended diatoms in the shallow waters (<60 cm) of the turbid tidal edge are described for single sites on two tidal flats–the Molenplaat in the Westerschelde estuary, and the Hond in the Ems-Dollard estuary, The Netherlands. High concentrations of chlorophyll-a (chl-a) were observed in the trailing edge of the ebbing tide in water depths of <20 cm, after which concentrations decreased markedly. Peak mean values were 19 μg chl-a l⁻¹ in 10 cm of water at the Molenplaat, and 45 μg chl-a l⁻¹ in 5 cm of water at the Hond. Similar trends were observed on the flooding tide, although peak values were far less pronounced (6 and 30 μg chl-a l⁻¹ respectively). Microscopic examination of the diatom community within the turbid tidal edge at the Molenplaat revealed that peaks in biomass were caused by suspended benthic diatoms, as well as the large centric diatom Coscinodiscus sp., particularly on the ebb tide. Planktonic diatoms other than Coscinodiscus sp. were more randomly distributed and did not appear to follow any particular trend. It would seem that as the tide recedes, resuspended benthic diatoms and large Coscinodiscus sp. cells become concentrated in the shallow water. However, the virtual absence of Coscinodiscus sp. from the leading edge of the flooding tide suggests that most of the resuspended cells do not settle to the seabed, but are washed away into the channels. The small peak of benthic diatoms at the leading edge of the flood tide is most likely resuspended locally from the sediment, along with large numbers of diatom frustules.     

隐虾类与海洋无脊椎动物的共栖关系（Ⅱ）

４隐虾类与腔肠动物的共栖关系隐虾类似乎对腔肠动物特别感兴趣，与之共栖者最多，达３０属，但绝大多数是与珊瑚虫纲（Ａｎｔｈｏｚｏａ），特别是石珊瑚目（Ｓｃｌｅｒａｃｔｉｎｉａ）、柳珊瑚目（Ｇｏｒｇｏｎａｃｅａ）和海葵目（Ａｃｔｉｎｉａｒｉａ）共栖，这与珊瑚虫纲动物固着生活的习性，是珊瑚礁构造的主体，其结构，特别是群体的结构可形成大量的小空间，极适于隐虾类的生活习性大有关系以下分别简述隐虾类与腔肠动物各类群的共栖关系。４．１珊瑚虫纲４．１．１石珊瑚目隐虾类共有１８属４２种已确定与石珊瑚共栖，其中滨虾属…     

Benthic foraminifers in upper Quaternary sediments of the southern Bering Sea: Distribution and paleoceanographic interpretations

Composition and distribution of benthic foraminifers, being coupled with isotopic-geochemical data on Upper Pleistocene and Holocene sediments from the southern Bering Sea (Core GC-11; 53°31′ N, 178°51′E, water depth 3060 m), demonstrate variations in bottom water properties during the last 54 kyr. Their abundance increased to some extent during short periods corresponding to warm Dansgaard-Oeshger interstadials 14, 12, 8, and 2 of marine isotopic stages (MIS) 3 and 2. The first and second deglaciation phases separated by the Younger Dryas cooling episode are marked by significant abundance peaks of benthic foraminifers (an order magnitude higher than in the glacial period), although their share in community of benthic and planktonic foraminifers taken together decreases. Species typical of stable high-productivity areas gain the dominant position. A significant proportion of agglutinated species in the Holocene sediments is indicative of Ca ions deficiency that accelerates dissolution of carbonate tests up to their disappearance approximately 2.5–3 ka ago.     

Colonization and evolution of lakes on the central Norwegian coast following deglaciation and land uplift 9500 to 7800 years B.P.

Invertebrate colonization of lakes following the uplift of land from the sea was studied in four lakes, currently situated between 39 and 24 m a.s.l., on the central Norwegian coast. The lakes were isolated from the sea between 9500 and 7700 years B.P. Animal and algal remains picked from core samples showed that the first colonizers preserved as fossils were usually members of the Chironomidae, Daphnidae/Chydoridae, Acarina, Porifera (Ephydatia mülleri and Spongilla lacustris), Bryozoa (Cristatella mucedo and Plumatella spp.) and Charophyta (Chara sp.). Of the chironomids, the genus Chironomus was present in the oldest lacustrine layers of all four lakes, but other genera recorded at the marine/lacustrine boundary were Dicrotendipes, Procladius (?), Einfeldia, Microtendipes, and Glyptotendipes. Remains of the caddis fly family Limnephilidae were also present in the earliest lacustrine sediments in Kvennavatnet and Kvernavatnet. The oldest invertebrate fauna is typical for mesotrophic lakes. However, chironomids and mites have been present in this area from at least about 10?500 years B.P. A diverse chironomid community was established between 300 and 800 years after isolation from the sea at Kvernavatnet on the island of Hitra, while only between 80 and 120 years passed before a comparably diverse community developed at Kvennavatnet on the mainland coast. A similar development of the invertebrate fauna occurred in Kvennavatnet, Kvernavatnet and Storkuvatnet. However, Litjvatnet deviates greatly from the ‘normal’ pattern because a tsunami disturbed the bottom sediments and fauna. The tsunami, a gigantic sea wave, was caused by a submarine slide from the Norwegian continental slope. It reached Litjvatnet, today located 24 m a.s.l., but was not traced in Storkuvatnet at 30 m a.s.l. This event happened about 7200 years B.P.     

Physical disturbance creates bacterial dominance of benthic biological communities in tropical deltaic environments of the Gulf of Papua

Unlike many reactive continental shelf mud deposits in temperate regions, bacteria and microfauna rather than macrofauna typically dominate benthic biomass and activities over large areas of the Gulf of Papua (GoP) deltaic complex, Papua New Guinea. During mid NW monsoon periods (Jan–Feb), macrofaunal densities at Gulf stations were relatively low (), large macroinfauna were absent (upper 25 cm), and small (), surface deposit-feeding polychaetes and tubiculous amphipods were dominant, reflecting a frequently destabilized seabed and high sedimentation/erosion rates. Although frequent physical disturbance generally inhibits development of macrobenthic communities, some regions of the Gulf deposits are periodically colonized and extensively bioturbated during quiescent periods, as shown by preserved biogenic sedimentary structures. Bacterial inventories integrated over the top 20 cm were extremely variable within each sub region of the clinoform complex. A possible bimodal pattern with bathymetric depth and distance offshore may occur: lowest-inventories within the sandy, proximal Fly River delta, an open Gulf inner topset zone (10–20 m) having sites of relatively high inventories, an open Gulf mid-topset region with intermediate values and less extreme variation, and the outer topset—upper foreset zone (40–50 m) where highest values are attained (). Various measures of microbial activity, including measures proportional to the cellular rRNA content and the proportion of dividing cells, indicate extremely productive populations over the upper 1-m of the seabed throughout the Gulf of Papua region. Bacterial biomass (0–20 cm) including data of Alongi et al. (1991, 1992, 1995) varied from a low of in intertidal mud banks to a high of in the topset—foreset zone. Macrofaunal biomass did not exceed in any sampled region, ranging from 0.009±0 to with no obvious correlation with bathymetric depth (1–63 m). Meiofaunal biomass was generally an order of magnitude lower than macrofaunal biomass. Relatively elevated bacterial biomass and high turnover rates are consistent with high measured rates of benthic remineralization, presumably reflecting the rapid response time of bacteria to physical reworking, the associated entrainment of organic substrate, and flushing of metabolites. Solute exchange is also enhanced below the directly mixed surface region, possibly producing ‘far field’ stimulation of microbes in underlying deposits. Physical reworking and reoxidation of sediments between 10 and 50 m water depth maintain suboxic, nonsulfidic conditions in the upper 0.5–1 m despite active microbial communities and high benthic remineralization rates.     

Untold muddy tales: Paleoenvironmental dynamics of a “barren” mudrock succession from a shallow Permian epeiric sea

During the late Paleozoic, the intracratonic Paraná Basin, Brazil, in central Gondwanaland, was covered by a huge (>1.600.000 km²), shallow and isolated epeiric sea. Within the Permian succession, oxygen-deficient facies are commonly recorded in the Mesosaurus-bearing Irati Formation (Cisuralian, Artinskian/Kungurian) and the overlaying Serra Alta Formation (Guadalupian, Wordian/Capitanian). Barren, dark-grey mudstones are the main facies preserved in this last unit, which has usually discouraged extensive and detailed stratigraphical and paleontological investigations. However, exhaustive sedimentological, taphonomic and paleontological surveys in those deposits reveal a dynamic and complex depositonal history. Based on sedimentary fabric, autochthonous to parautochthonous occurrences of shelly benthic invertebrates (bivalves) and the presence/absence of concretion-bearing and phosphate-rich layers, we report variations in the oxygen levels of bottom and pore waters, in bathymetry, sedimentation rates, and changes in benthic colonization. Our data indicate that the deposition of this “apparently barren” mudstone-dominated succession was driven by a complex interplay of variations in sedimentation rate and oxygen pulses tied to tectonic and climate changes. Three distinct populations or invertebrate paleocommunities were recorded, which were adapted to (a) normal background low-oxygen (dysoxic) conditions (i.e., minute infaunal suspension-feeding bivalves associated with the trace fossil Planolites), (b) chemically toxic (anoxic/extreme dysoxic) substrates, including gigantic burrowing bivalves (probable chemosymbiotic taxa), and (c) oxic/dysoxic substrates following short-term bottom disruptions.