Primary production by benthic microalgae in nearshore marine sediments of Signy Island,Antarctica

Summary During the austral summer of 1987/1988, three 24 h in situ primary productivity measurements were made at a nearshore sublittoral site on the east coast of Signy Island, Antarctica. The first experiment in December, coincided with the peak of the benthic algal bloom as shown by benthic chlorophyll measurements and a primary productivity rate of 700.9 mg carbon m^–2 day^–1. In January, the experiment was undertaken during the peak of the phytoplankton bloom when light intensities reaching the benthos were greatly reduced. A rate of 313.4 mg carbon m^–2 day^–1 was measured, half that of the previous month. In March the phytoplankton bloom had died off, benthic light intensities had increased and production was 391.8 mg m^–2 day^–1. The experiments indicate changes in benthic microalgal activity during the summer, linked to changes in the benthic light climate. Compared with previous measurements of phytoplanktonic activity at Signy, the microphytobenthos seems to be an important source of primary production. A production estimate of 100.9 mg carbon m^–2, for the ice-free summer period, lies within the range of values of results from other polar studies.     

Seasonal Dynamics of Benthic and Planktonic Algae in a Nutrient‐Rich Lowland River (Spree,Germany)

We studied chlorophyll a (chl. a), biovolume and species composition of benthic algae and phytoplankton in the eutrophic lower River Spree in 1996. The chl. a concentration was estimated as 3.5 (2.7–4.5) µg/cm² for epipsammon, 9.4 (7.4–11.9) µg/cm² for epipelon and 6.7 (5.7–7.8) µg/cm² for the epilithon (median and 95% C. L.). The mean total biomass of benthic algae was significantly higher (6.0 µg chl. a/cm²) than the areal chl. a content of the pelagic zone (1.6 µg chl. a/cm²). Although certain phytoplankton taxa were abundant in the periphyton, benthic taxa generally dominated the assemblages. Seasonal dynamics of benthic algae were probably controlled by light and nitrate supply (sand), discharge fluctuations (sand, mud) and invertebrate grazing (stones). This paper shows the importance of benthic algae even in phytoplankton‐rich lowland rivers with sandy or muddy sediments. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Instantaneous benthic response to different organic matter quality: In situ experiments in the Benguela Upwelling System

The benthic community in continental slope and deep-sea sediments of the Benguela Upwelling System was supplied with ¹³C-labelled organic matter (OM) of two different qualities using a benthic chamber lander. Freeze-dried cultures of Skeletonema costatum served as 'fresh' OM. 'Altered' OM of the same material had been additionally dialysed to remove low-molecular weight compounds. In order to investigate the benthic response pattern, mineralization of labelled OM, uptake by macrofauna and incorporation into bacteria were followed over 18-36 h. Total oxygen uptake was not affected beyond natural variation by the OM addition. Mineralization dominated the ¹³C-labelled phytodetritus processing, constituting 71-95% of the total processed OM. Bacterial incorporation of phytodetrital carbon exceeded macrofaunal uptake at all stations. Stations situated in a major centre of OM deposition showed phytodetritus processing rates on average twice as high as outside the depocentre. Phytodetritus processing was 1.5, 2.5 and 4.3 times higher for fresh than for altered OM at 605, 1019 and 1335 m water depth, respectively. Our observations clearly indicate the importance of OM quality on mineralization rates.     

Biodegradation of photosynthetically produced extracellular organic carbon from intertidal benthic algae

14C-labeled extracellular products of a natural microphytobenthic community and two species of benthic diatoms (Nitzschia hybridaeformis and Amphora coffeaeformis) were fractionated into extracellular dissolved organic carbon (14C-EDOC), organic carbon extracted with EDTA (14C-EDTA-extractable OC) and extracellular polymeric substances (14C-EPS). The biodegradation of this labeled extracellular organic carbon by bacteria in sediments was examined to determine the processes of enzymatic degradation of photosynthetically-produced extracellular organic carbon from microphytobenthos in an intertidal flat ecosystem. In addition, primary production as well as extracellular enzyme activities (beta- and alpha-glucosidase) were measured to evaluate the possible relationship between organic carbon production and microbiological degradation at the Isshiki intertidal flat in Mikawa Bay, Japan. With all three 14C-fractions extracted from a natural microphytobenthic assemblage and two species of benthic diatoms, more than 50% of the added substrates were mineralized within 24 h by the bacterial community in sediments. At that time, the percentage of high-molecular-weight compounds (>5 K MW) to total MW compounds of 14C-EDTA-extractable OC and 14C-EPS fractions decreased within 24 h from 50.9 to 6.6% and 74.5 to 11.1%, respectively. In situ, beta- and alpha-glucosidase activity in sediment was higher than in the seawater column (at a depth of 1 m), though the photosynthetic production of microphytobenthos was equal to that of phytoplankton. Based on our previous studies that microphytobenthos produced much more extracellular products than phytoplankton, it is assumed from these results that carbon flowing into the microbial loop through the mediation of enzymatic degradation of extracellular products in a benthic system exceeds that in the overlying water column.     

Planktonic bacterial DNA and RNA synthesis from algal extracellular products in a eutrophic lake

Abstract In order to better understand bacterial growth in the natural environment, a tracer experiment was carried out by adding [¹⁴C]bicarbonate to samples from a eutrophic lake and enclosures which were constructed inside the lake to provide different substrate conditions. It was shown that extracellular dissolved organic carbon (EDOC) released from photosynthetic phytoplankton was one of the most important carbon sources for the growth of planktonic bacteria in the aquatic environment studied here. About 5% of photosynthetically fixed ¹⁴C was transferred to the bacterial fraction between 4 and 8 h after the addtion of the [¹⁴C]bicarbonate. This increased to 10% during the following 16 h in the dark. Between 50% and 60% of bacterial ¹⁴C from EDOC was transformed to bacterial macromolecules. Analysis of cellular components showed that 36–39% of macromolecules produced were DNA during the light incubation, although the proportion of DNA relative to total macromolecules decreased in the dark. RNA synthesis increased with time, from 25% to 38% during the dark incubation. There was a correlation between bacterial growth rates and the ratio of RNA/DNA synthesized from EDOC (r = 0.81). However, the RNA/DNA ratio was very small, 1.8 for all samples examined, compared with previous data obtained for the laboratory bacteria. This is probably due to the very slow growth rate of the natural planktonic bacteria.     

A small population of planktonic Flavobacteria with disproportionally high growth during the spring phytoplankton bloom in a prealpine lake

Bacterioplankton growth in temperate Lake Zurich (Switzerland) was studied during the spring phytoplankton bloom by in situ techniques and short-term dilution bioassays. A peak of chlorophyll a (Chl a ) concentrations was followed by a rise of bacterial cell numbers and leucine assimilation rates, of the proportions of cells incorporating 5-bromo-2-deoxyuridine (BrdU), and of community net growth rates in dilution cultures. Incorporation of BrdU was low in Betaproteobacteria (2 ± 1%), indicating that these bacteria did not incorporate the tracer. Pronounced growth of Betaproteobacteria in the enrichments was only observed after the decline of the phytoplankton bloom. An initial peak in the proportions of BrdU-positive Actinobacteria (30%) preceded a distinct rise of their cell numbers during the period of the Chl a maximum. Cytophaga–Flavobacteria (CF) changed little in numbers, but featured high proportions of BrdU-positive cells (28 ± 12%). Moreover, CF represented > 90% of all newly formed cells in dilution cultures before and during the phytoplankton bloom. One phylogenetic lineage of cultivable Flavobacteria (FLAV2) represented a small (0.5–1%) but highly active population in lake plankton. The growth rates of FLAV2 in dilution cultures doubled during the period of the Chl a maximum, indicating stimulation by phytoplankton exudates. Thus, CF, and specifically Flavobacteria , appeared to be substantially more important for carbon transfer in Lake Zurich spring bacterioplankton than was suggested by their standing stocks. The high in situ growth potential of these bacteria might have been counterbalanced by top-down control.     

Impact of warming on phyto‐bacterioplankton coupling and bacterial community composition in experimental mesocosms

Global warming is assumed to alter the trophic interactions and carbon flow patterns of aquatic food webs. The impact of temperature on phyto‐bacterioplankton coupling and bacterial community composition (BCC) was the focus of the present study, in which an indoor mesocosm experiment with natural plankton communities from the western Baltic Sea was conducted. A 6°C increase in water temperature resulted, as predicted, in tighter coupling between the diatom‐dominated phytoplankton and heterotrophic bacteria, accompanied by a strong increase in carbon flow into bacterioplankton during the phytoplankton bloom phase. Suppressed bacterial development at cold in situ temperatures probably reflected lowered bacterial production and grazing by protists, as the latter were less affected by low temperatures. BCC was strongly influenced by the phytoplankton bloom stage and to a lesser extent by temperature. Under both temperature regimes, Gammaproteobacteria clearly dominated during the phytoplankton peak, with Glaciecola sp. as the single most abundant taxon. However, warming induced the appearance of additional bacterial taxa belonging to Betaproteobacteria and Bacteroidetes. Our results show that warming during an early phytoplankton bloom causes a shift towards a more heterotrophic system, with the appearance of new bacterial taxa suggesting a potential for utilization of a broader substrate spectrum.     

毛乌素沙地固沙林发育过程中土壤有机碳库稳定性特征

为揭示固沙林发育过程中土壤固碳效应与有机碳库稳定性特征,基于时空替代法,采集陕北毛乌素沙地治沙区从半固定沙地到林龄分别为22、32、53年乔木和灌木林地表层土壤,分析了土壤总有机碳、氧化活性碳和惰性有机碳含量,以及矿化碳排放、分解比例的变异特征.结果表明:相比半固定沙地,22~53年灌木和乔木林土壤总有机碳增幅来自惰性碳,分别为3.5~6.2和3.2~7.7 g·kg^-1,来自氧化活性碳的分别为2.8~3.4和1.3~2.8 g·kg^-1,并且灌木和乔木林土壤氧化活性碳所占比例未发生显著变化,分别维持在37.0%和26.8%,但土样经恒温培养60 d后,该比例分别下降至25.7%和17.4%.培养结束时,2种林地22~53年土壤有机碳矿化率无显著差异;所有固沙林样地矿化碳的损失有76.9%~98.7%来自于氧化活性碳,仅1.3%~23.5%来自于惰性碳.与最高碳分解速率相比,土壤碳累积释放量与β-葡萄糖苷酶、脱氢酶活性的相关性更强,但酶活性在32~53年林地无显著差异.综上,随着固沙林林龄增加,土壤有机碳趋向少排多存碳的稳定性特征,乔木固沙林固碳效果优于灌木固沙林.     

Qualitative importance of the microbial loop and plankton community structure in a eutrophic lake during a bloom of cyanobacteria

Plankton community structure and major pools and fluxes of carbon were observed before and after culmination of a bloom of cyanobacteria in eutrophic Frederiksborg Slotssø, Denmark. Biomass changes of heterotrophic nanoflagellates, ciliates, microzooplankton (50 to 140 μm), and macrozooplankton (larger than 140 μm) were compared to phytoplankton and bacterial production as well as micro- and macrozooplankton ingestion rates of phytoplankton and bacteria. The carbon budget was used as a means to examine causal relationships in the plankton community. Phytoplankton biomass decreased and algae smaller than 20 μm replacedAphanizomenon after the culmination of cyanobacteria. Bacterial net production peaked shortly after the culmination of the bloom (510 μg C liter^?1 d^?1 and decreased thereafter to a level of approximately 124 μg C liter^?1 d^?1. Phytoplankton extracellular release of organic carbon accounted for only 4–9% of bacterial carbon demand. Cyclopoid copepods and small-sized cladocerans started to grow after the culmination, but food limitation probably controlled the biomass after the collapse of the bloom. Grazing of micro- and macrozooplankton were estimated from in situ experiments using labeled bacteria and algae. Macrozooplankton grazed 22% of bacterial net production during the bloom and 86% after the bloom, while microzooplankton (nauplii, rotifers and ciliates larger than 50 μm) ingested low amounts of bacteria and removed 10–16% of bacterial carbon. Both macro-and microzooplankton grazed algae smaller than 20 μm, although they did not control algal biomass. From calculated clearance rates it was found that heterotrophic nanoflagellates (40–440 ml^?1) grazed 3–4% of the bacterial production, while ciliates smaller than 50 μm removed 19–39% of bacterial production, supporting the idea that ciliates are an important link between bacteria and higher trophic levels. During and after the bloom ofAphanizomenon, major fluxes of carbon between bacteria, ciliates and crustaceans were observed, and heterotrophic nanoflagellates played a minor role in the pelagic food web.     

The importance of bacterial utilization of released phytoplankton photosynthate in two humic forest lakes in southern Finland

Bacterial utilization of photosynthetically fixed dissolved organic carbon (PDOC) released from natural phytoplankton assemblages was studied in two small, extremely humic, forest lakes in southern Finland. Bacterial activity (measured as uptake of ¹⁴C-glucose) and phytoplankton photosynthesis (measured as light uptake of ¹⁴CO₂) could be most effectively separated using Nuclepore filters of pore size 1–2 μm. Released PDOC was 10–67% of total phytoplankton carbon fixation during in situ experiments, and represented about 0.1% of total DOC. Net uptake of PDOC by bacteria was found to be about 20% during 24 hour laboratory incubations, although about 40% of PDOC present at the start of an experiment could be utilized by bacteria during a 24 hour period. PDOC does not provide a quantitatively important substrate supply for bacterial respiration in humic forest lakes.     

Oxygen dynamics in shelf seas sediments incorporating seasonal variability

Shelf sediments play a vital role in global biogeochemical cycling and are particularly important areas of oxygen consumption and carbon mineralisation. Total benthic oxygen uptake, the sum of diffusive and faunal mediated uptake, is a robust proxy to quantify carbon mineralisation. However, oxygen uptake rates are dynamic, due to the diagenetic processes within the sediment, and can be spatially and temporally variable. Four benthic sites in the Celtic Sea, encompassing gradients of cohesive to permeable sediments, were sampled over four cruises to capture seasonal and spatial changes in oxygen dynamics. Total oxygen uptake (TOU) rates were measured through a suite of incubation experiments and oxygen microelectrode profiles were taken across all four benthic sites to provide the oxygen penetration depth and diffusive oxygen uptake (DOU) rates. The difference between TOU and DOU allowed for quantification of the fauna mediated oxygen uptake and diffusive uptake. High resolution measurements showed clear seasonal and spatial trends, with higher oxygen uptake rates measured in cohesive sediments compared to the permeable sediment. The significant differences in oxygen dynamics between the sediment types were consistent between seasons, with increasing oxygen consumption during and after the phytoplankton bloom. Carbon mineralisation in shelf sediments is strongly influenced by sediment type and seasonality.     

The submicron size fraction of seawater containing high numbers of virus particles as bioactive agent in unicellular plankton community successions

The effect of moderately (2.5 times) increasing the concentrationof the submicron size fraction (2–200 nm) of seawateron unicellular heterotrophic plankton organisms was tested ina series of experiments during algal bloom conditions. Usingelectron microscopy, vims-like particles (VLPs) were by farthe most visible component in this size fraction. We incubatedseawater. which was collected in late winter, in slowly rotatingcylindrical tanks that were exposed to spring temperature andlight conditions. This led to the onset of a mixed diatom bloomwhich was delayed by several days in submicron size-enrichedtreatments. Later, phytoplankton biomass reached values evenslightly higher than in unaltered incubations. The abundanceof heterotrophic unicellular plankton and VLPs exhibited morepronounced oscillation amplitudes during the early phase ofsuccession, thus being stimulated by additional submicron-sizedmaterial (including virions). VLPs are suggested as a possiblesupplementary diet for heterotrophic flagellates. From 190 honwards, toward the end of the incubation experiments (550 h).heterotrophic flagellate density was no longer influenced byinitial enrichment with the submicron size fraction. In contrast,bacteria were consistently repressed in the last two-thirdsof the experimental duration. Our results support the existenceof bioactive substances in the 2-200 nm size fraction, mostprobably virus particles, which are capable of influencing thedynamics and oscillation patterns in plankton community successionaccording to their concentration during algal bloom events.     

NUTRIENT LIMITATION OF BENTHIC ALGAE IN LAKE MICHIGAN: THE ROLE OF SILICA1

In the Laurentian Great Lakes, phytoplankton growth and biomass are secondarily limited by silica (Si), as a result of phosphorus (P) enrichment. Even modest levels of P enrichment can induce secondary Silimitation, which, in turn, promotes a shift from the native diatom phytoplankton flora to chlorophyte and cyanobacteria species. However, very little is known about the nutritional status of benthic populations and their response to nutrient enrichment. Two experiments were performed in the littoral zone of Lake Michigan where nutrients were delivered to in situ benthic algal (episammic and epilithic) assemblages using nutrient‐diffusing substrata. In order to test the hypothesis that benthic algae in Lake Michigan are Si limited, a 2 × 3 factorial experiment was used to deliver all combinations of Si, N, and P to resident assemblages growing on artificial substrata composed of natural (Si rich) versus calcium carbonate (Si poor) sand. A second experiment utilized a serial enrichment to evaluate the role of Si in mediating changes in taxonomic composition. These findings indicate that benthic algae in Lake Michigan exhibit signs of secondary Si limitation, and that their response to enrichment is similar to the phytoplankton. Moreover, natural sand substrata may provide a source of Si to resident benthic algae.     

Biochemical characteristics of surface sediments on the eastern Weddell Sea continental shelf, Antarctica: is there any evidence of seasonal patterns?

Biochemical characteristics of seafloor sediment off Austasen in the southeastern Weddell Sea were assayed in samples recovered in the early autumn and late spring of 2000 and 2003, respectively. Sediment was separated in the grain-size fractions >200 μm and <200 μm to distinguish biochemical characteristics in the fraction available for benthic suspension feeders (<200 μm). In the bulk sediment, the lipid (LPD) and carbohydrate (CHO) contents were significantly different between seasons with higher LPD content in the early autumn and higher CHO content in the late spring. In the grain-size fractions <200 μm, the LPD and protein (PRT) contents were significantly higher in the early autumn meaning that in this season the fraction available for benthic suspension feeders presented higher nutritive value. The relatively higher CHO concentrations observed in each fraction in the late spring were attributed to refractory matter, whereas the higher PRT and LPD concentrations found during the early autumn were associated with planktonic material settled after the summer phytoplankton bloom. Our results suggest that there is seasonal variation in the composition of organic matter in the sediment, with better nutritive quality in the early autumn, especially in the grain-size fraction available for benthic suspension feeders. These variations also suggest that the benthic community exploits the fresh organic matter accumulated after the summer throughout the Antarctic dark months leaving the sediment almost exhaust of LPD and with higher CHO contents, presumably of refractory nature, at the onset of the seasonal phytoplankton bloom of the following year.     

Rates of Benthic Protozoan Grazing on Free and Attached Sediment Bacteria Measured with Fluorescently Stained Sediment

In order to determine the importance of benthic protozoa as consumers of bacteria, grazing rates have been measured by using monodispersed fluorescently labeled bacteria (FLB). However, high percentages of nongrazing benthic protists are reported in the literature. These are related to serious problems of the monodispersed FLB method. We describe a new method using 5-(4,6-dichlorotriazin-2-yl)-aminofluorescein (DTAF)-stained sediment to measure in situ bacterivory by benthic protists. This method is compared with the monodispersed FLB technique. Our estimates of benthic bacterivory range from 61 to 73 bacteria protist^-1 h^-1 and are about twofold higher than the results of the monodispersed FLB method. The number of nongrazing protists after incubation for 15 min with DTAF-stained sediment is in agreement with theoretical expectation. We also tested the relative affinity for FLB of protists and discuss the results with respect to a grazing model.     

Estimating Bacterioplankton Production by Measuring [H]thymidine Incorporation in a Eutrophic Swedish Lake

Bacterioplankton abundance, [H]thymidine incorporation, CO(2) uptake in the dark, and fractionated primary production were measured on several occasions between June and August 1982 in eutrophic Lake Norrviken, Sweden. Bacterioplankton abundance and carbon biomass ranged from 0.5 x 10 to 2.4 x 10 cells liter and 7 to 47 mug of C liter, respectively. The average bacterial cell volume was 0.185 mum. [H]thymidine incorporation into cold-trichloroacetic acid-insoluble material ranged from 12 x 10 to 200 x 10 mol liter h. Bacterial carbon production rates were estimated to be 0.2 to 7.1 mug of C liter h. Bacterial production estimates from [H]thymidine incorporation and CO(2) uptake in the dark agreed when activity was high but diverged when activity was low and when blue-green algae (cyanobacteria) dominated the phytoplankton. Size fractionation indicated negligible uptake of [H]thymidine in the >3-mum fraction during a chrysophycean bloom in early June. We found that >50% of the H activity was in the >3-mum fraction in late August; this phenomenon was most likely due to Microcystis spp., their associated bacteria, or both. Over 60% of the CO(2) uptake in the dark was attributed to algae on each sampling occasion. Algal exudate was an important carbon source for planktonic bacteria. Bacterial production was roughly 50% of primary production.     

Correlations between phytoplankton,organic detritus and carbon in North Sea waters during the Fladenground Experiment (FLEX' 76)

Quantitative analysis of phytoplankton, organic detritus and dissolved organic carbon was carried out, during the spring bloom, in a specific North Sea area (58° 55 N, O° 32 E) in the course of an international and multidisciplinary marine investigation (FLEX' 76). An attempt was made to correlate the fluctuations of these variables; the analysis, however, was complicated by massive water movements. A maximum of the phytoplankton bloom coincided with a maximum of organic detritus, while both variables turned out to be inversely correlated with the dissolved organic carbon content. The initial, high amount of dissolved organic carbon declined at a daily rate of ca. 13% during a period of non-interference by water movements. Two thirds of the dissolved organic carbon content are assumed to be transformed into particulate organic matter whilst the rest disappears through still unknown channels.     

Linkages between bacterioplankton community composition, heterotrophic carbon cycling and environmental conditions in a highly dynamic coastal ecosystem

We used mesocosm experiments to study the bacterioplankton community in a highly dynamic coastal ecosystem during four contrasting periods of the seasonal cycle: winter mixing, spring phytoplankton bloom, summer stratification and autumn upwelling. A correlation approach was used in order to measure the degree of coupling between the dynamics of major bacterial groups, heterotrophic carbon cycling and environmental factors. We used catalysed reporter deposition-fluorescence in situ hybridization to follow changes in the relative abundance of the most abundant groups of bacteria (Alphaproteobacteria, Gammaproteobacteria and Bacteroidetes). Bacterial carbon flux-related variables included bacterial standing stock, bacterial production and microbial respiration. The environmental factors included both, biotic variables such as chlorophyll-a concentration, primary production, phytoplankton extracellular release, and abiotic variables such as the concentration of dissolved inorganic and organic nutrients. Rapid shifts in the dominant bacterial groups occurred associated to environmental changes and bacterial bulk functions. An alternation between Alphaproteobacteria and Bacteroidetes was observed associated to different phytoplankton growth phases. The dominance of the group Bacteroidetes was related to high bacterial biomass and production. We found a significant, non-spurious, linkage between the relative abundances of major bacterial groups and bacterial carbon cycling. Our results suggest that bacteria belonging to these major groups could actually share a function in planktonic ecosystems.     

Identification of DNA-synthesizing bacterial cells in coastal North Sea plankton

We describe a method for microscopic identification of DNA-synthesizing cells in bacterioplankton samples. After incubation with the halogenated thymidine analogue bromodeoxyuridine (BrdU), environmental bacteria were identified by fluorescence in situ hybridization (FISH) with horseradish peroxidase (HRP)-linked oligonucleotide probes. Tyramide signal amplification was used to preserve the FISH staining during the subsequent immunocytochemical detection of BrdU incorporation. DNA-synthesizing cells were visualized by means of an HRP-labeled antibody Fab fragment and a second tyramide signal amplification step. We applied our protocol to samples of prefiltered (pore size, 1.2 micro m) North Sea surface water collected during early autumn. After 4 h of incubation, BrdU incorporation was detected in 3% of all bacterial cells. Within 20 h the detectable DNA-synthesizing fraction increased to >14%. During this period, the cell numbers of members of the Roseobacter lineage remained constant, but the fraction of BrdU-incorporating Roseobacter sp. cells doubled, from 24 to 42%. In Alteromonas sp. high BrdU labeling rates after 4 to 8 h were followed by a 10-fold increase in abundance. Rapid BrdU incorporation was also observed in members of the SAR86 lineage. After 4 h of incubation, cells affiliated with this clade constituted 8% of the total bacteria but almost 50% of the visibly DNA-synthesizing bacterial fraction. Thus, this clade might be an important contributor to total bacterioplankton activity in coastal North Sea water during periods of low phytoplankton primary production. The small size and low ribosome content of SAR86 cells are probably not indications of inactivity or dormancy.