Quantitative Importance,Composition, and Seasonal Dynamics of Protozoan Communities in Polyhaline versus Freshwater Intertidal Sediments

The quantitative importance and composition of protozoan communities was investigated in sandy and silty intertidal sediments of a polyhaline and a freshwater site in the Schelde estuary. Total biomass of the protozoans studied, integrated over the upper 4 cm of the sediment, ranged from 41 to 597 mg C m^–2 and was in the same order of magnitude at the polyhaline and the freshwater intertidal site. Nanoheterotrophs were the dominant protozoans, in terms of both abundance and biomass. Ciliate abundances appeared to be largely determined by physical constraints, namely, the amount of interstitial space and hydrodynamic disturbances. It remains unclear which factors control nanoheterotrophic abundances and biomasses, which showed comparatively little seasonal and between-site fluctuations. Salinity differences were clearly reflected in the protozoan community composition. The dominant role of sessile ciliates is a unique feature of sediments in the freshwater tidal reaches, which can be attributed to the dynamic nature of sedimentation and resuspension processes associated with the maximum turbidity zone. Based on biomass ratios and estimated weight-specific metabolic rates, protozoa possibly accounted for ~29 to 96% of the estimated combined metabolic rate of protozoan and metazoan consumers at our sampling stations in late spring/early autumn. The contribution of protozoa to this combined metabolic rate was higher at the sandy than at the silty stations and was mainly accounted for by the nanoheterotrophs. These data emphasize the potential importance of small protozoa in sediments and suggest that protozoa are important components of benthic food webs.     

Impact of Water Column Acidification on Protozoan Bacterivory at the Lake Sediment-Water Interface

Although the impact of acidification on planktonic grazer food webs has been extensively studied, little is known about microbial food webs either in the water column or in the sediments. Protozoon-bacterium interactions were investigated in a chronically acidified (acid mine drainage) portion of a lake in Virginia. We determined the distribution, abundance, apparent specific grazing rate, and growth rate of protozoa over a pH range of 3.6 to 6.5. Protozoan abundance was lower at the most acidified site, while abundance, in general, was high compared with other systems. Specific grazing rates were uncorrelated with pH and ranged between 0.02 and 0.23 h^-1, values similar to those in unacidified systems. The protozoan community from an acidified station was not better adapted (P = 0.95) to low-pH conditions than a community from an unacidified site (multivariate analysis of variance on growth rates for each community incubated at pHs 4, 5, and 6). Both communities had significantly lower (P < 0.05) growth rates at pHs 4 and 5 than at pH 6. Reduced protozoan growth rates coupled with high grazing rates and relatively higher bacterial yields (ratio of bacterial-protozoan standing stock) at low pH indicate reduced net protozoan growth efficiency and a metabolic cost of acidification to the protozoan community. However, the presence of an abundant, neutrophilic protozoan community and high bacterial grazing rates indicates that acidification of Lake Anna has not inhibited the bacterium-protozoon link of the sediment microbial food web.     

Regulation of oxidative phosphorylation complex activity: effects of tissue-specific metabolic stress within an allometric series and acute changes in workload

The concentration of mitochondrial oxidative phosphorylation complexes (MOPCs) is tuned to the maximum energy conversion requirements of a given tissue; however, whether the activity of MOPCs is altered in response to acute changes in energy conversion demand is unclear. We hypothesized that MOPCs activity is modulated by tissue metabolic stress to maintain the energy-metabolism homeostasis. Metabolic stress was defined as the observed energy conversion rate/maximum energy conversion rate. The maximum energy conversion rate was assumed to be proportional to the concentration of MOPCs, as determined with optical spectroscopy, gel electrophoresis, and mass spectrometry. The resting metabolic stress of the heart and liver across the range of resting metabolic rates within an allometric series (mouse, rabbit, and pig) was determined from MPOCs content and literature respiratory values. The metabolic stress of the liver was high and nearly constant across the allometric series due to the proportional increase in MOPCs content with resting metabolic rate. In contrast, the MOPCs content of the heart was essentially constant in the allometric series, resulting in an increasing metabolic stress with decreasing animal size. The MOPCs activity was determined in native gels, with an emphasis on Complex V. Extracted MOPCs enzyme activity was proportional to resting metabolic stress across tissues and species. Complex V activity was also shown to be acutely modulated by changes in metabolic stress in the heart, in vivo and in vitro. The modulation of extracted MOPCs activity suggests that persistent posttranslational modifications (PTMs) alter MOPCs activity both chronically and acutely, specifically in the heart. Protein phosphorylation of Complex V was correlated with activity inhibition under several conditions, suggesting that protein phosphorylation may contribute to activity modulation with energy metabolic stress. These data are consistent with the notion that metabolic stress modulates MOPCs activity in the heart.     

An Under-appreciated Component of Biodiversity in Plankton Communities: The Role of Protozoa in Lake Michigan (A Case Study)

Recent technological advances have led to the discovery that free-living, planktonic protozoa are ubiquitous in nature and appear to be important components of pelagic food webs (e.g., fluorescent straining, flow cytometry). Despite this, limited information exists tying their seasonality to rate processes that drive succession patterns. The abundance, and seasonal growth and grazing loss of an entire protozoan assemblage were evaluated in Lake Michigan. The protozoan assemblage was species-rich (100 taxa) and abundant throughout the year in Lake Michigan. Nano-sized protozoa (Hnano and Pnano, <20 μm in size) ranged in abundance from 10² to 10³ cells ml⁻¹, while micro-protozoa (Hmicro and Pmico, >20 and <200 μm in size) ranged in abundance from 4 to 17 cells ml⁻¹. The biomass of Hnano and Hmicro by itself represented more than 70–80% of crustacean zooplankton biomass, while Pnano and Pmicro constituted nearly 50% of phytoplankton biomass. Protozoa exhibited growth rates comparable to other components of the plankton in Lake Michigan, and some populations grew at rates similar to maximum rates determined in the laboratory (rates of 1–2 day⁻¹). Overall, it appears that macro-zooplankton predation is a major loss factor counter-balancing growth with only small differences between the two rate processes (<0.1 day⁻¹). Discrepancies between growth and grazing loss in the spring were likely attributed to sedimentation losses for larger species of tintinnids and dinoflagellates (Codonella, Tintinnidium, and Gymnodinium) that can account for their occurrence in the deep chlorophyll layer. In the summer, carnivory among similar sized species (Chromulina and small ciliates) may be additional loss factors impinging on the protozoan assemblage.     

A high standard metabolic rate constrains juvenile growth

The allocation of energy to various components of an individual's energy budget is often viewed as a competitive process. As such, a tradeoff may exist between production (growth) and maintenance metabolism. One view of a potential tradeoff, termed “the principle of allocation”, suggests that individuals with lower maintenance metabolic expenditures may have higher growth rates. To determine whether such a tradeoff exists, I analyzed the relationship between growth rate and maintenance metabolism of 225 juvenile snapping turtles housed in the laboratory. I measured growth from hatching to 6 months of age, and then measured oxygen consumption and calculated standard metabolic rate. Mean growth rate was 0.19 g d⁻ and mean standard metabolic rate (SMR) was 1.41 kJ d⁻. Maintenance metabolism and growth were negatively correlated after both were adjusted for body mass. The results support the “principle of allocation” theory: individuals with higher standard metabolic rates tended to have low growth rates.     

The Warburg effect and mitochondrial oxidative phosphorylation: Friends or foes?

Cancer cells display an altered energy metabolism, which was proposed to be the root of cancer. This early discovery was done by O. Warburg who conducted one of the first studies of tumor cell energy metabolism. Taking advantage of cancer cells that exhibited various growth rates, he showed that cancer cells display a decreased respiration and an increased glycolysis proportional to the increase in their growth rate, suggesting that they mainly depend on fermentative metabolism for ATP generation.Warburg's results and hypothesis generated controversies that are persistent to this day. It is thus of great importance to understand the mechanisms by which cancer cells can reversibly regulate the two pathways of their energy metabolism as well as the functioning of this metabolism in cell proliferation. In this review, we discuss of the origin of the decrease in cell respiratory rate, whether the Warburg effect is mandatory for an increased cell proliferation rate, the consequences of this effect on two major players of cell energy metabolism that are ATP and NADH, and the role of the microenvironment in the regulation of cellular respiration and metabolism both in cancer cell and in yeast.     

Continuous culture of the ciliate Tetrahymena pyriformis on Escherichia coli

The ciliated protozoan Tetrahymena pyriformis was grown in a chemostat fed with a culture of Escherichia coli overflowing from another chemostat. Densities of the protozoan and bacterial populations, mean volume of protozoan cells, yields of protozoan volumes and numbers, and filtering rates of protozoans per cell and per unit volume of biomaterial were determined at five different dilution rates. The data obtained supplement other data already available for the popular test organism T. pyriformis, and they are also comparable with data available for related ciliates.     

中国淡水原生动物多样性及其所受威胁

概述了地球上原生动物多样性现状,指出它在生物学界中的地位及其对人类的作用。从生态学概念出发,把淡水原生动物作为一个集合类群,对其多样性和所受威胁进行了讨论。以武汉东湖为例,讨论了半个世纪来由于人类活动的影响引起湖泊富营养化后,原生动物丰度增长了10倍,优势种出现替代现象,并有明显的小型化倾向。以长江三峡地区为例,分析了工业污染能损伤70％原生动物种类。原生动物多样性指数变化表明：在有机污染影响下作为饮用水的汉江其水质已达临界边缘。对待开发的西南武陵山地区生物资源考察中,用原生动物多样性现状阐明该地区水生态系的稳定性。     

Effects of body size and temperature on population growth

For at least 200 years, since the time of Malthus, population growth has been recognized as providing a critical link between the performance of individual organisms and the ecology and evolution of species. We present a theory that shows how the intrinsic rate of exponential population growth, rmax, and the carrying capacity, K, depend on individual metabolic rate and resource supply rate. To do this, we construct equations for the metabolic rates of entire populations by summing over individuals, and then we combine these population-level equations with Malthusian growth. Thus, the theory makes explicit the relationship between rates of resource supply in the environment and rates of production of new biomass and individuals. These individual-level and population-level processes are inextricably linked because metabolism sets both the demand for environmental resources and the resource allocation to survival, growth, and reproduction. We use the theory to make explicit how and why rmax exhibits its characteristic dependence on body size and temperature. Data for aerobic eukaryotes, including algae, protists, insects, zooplankton, fishes, and mammals, support these predicted scalings for rmax. The metabolic flux of energy and materials also dictates that the carrying capacity or equilibrium density of populations should decrease with increasing body size and increasing temperature. Finally, we argue that body mass and body temperature, through their effects on metabolic rate, can explain most of the variation in fecundity and mortality rates. Data for marine fishes in the field support these predictions for instantaneous rates of mortality. This theory links the rates of metabolism and resource use of individuals to life-history attributes and population dynamics for a broad assortment of organisms, from unicellular organisms to mammals.     

Functional Groups in the Protozoa: Roles in Differing Ecosystems1,2

Feeding habits of freshwater protozoa were used to group species into functional, trophic groups. Community structure in differing ecosystems was examined in relation to the number of species occurring in the functional group categories. Six wetland ecosystems and a large river ecosystem were studied. Changes in community structure during the colonization of artificial substrates were also examined. Changes during colonization were studied in a mesotrophic lake, in low-order streams, and in laboratory microecosystems. In the latter case, the response of colonizing communities to a heavy metal toxicant was studied. All communities studied were dominated by bactivorous-detritivorous species and, to a lesser extent, by photosynthetic species. The chief functional role of substrate-associated protozoans appears to be the processing of dead organic matter and its associated bacterial flora. Functional groups utilizing resources other than detrital or mineral nutrients (saprotrophs, algivores, omnivores, and predators) were always minor community components. Colonizing communities were often dominated by photosynthetic species during early colonization stages but were again dominated by bactivorous-detritivorous species at species equilibrium. Low levels of toxicant (Cd) reduced numbers of both photosynthetic and bactivorous-detritivorous species. Higher toxicant levels virtually eliminated photosynthetic species and reduced bacterial detritivores by over one-half. Roles of protozoan species in ecosystems are closely tied to the processing of detritus and the recycling of mineral nutrients. Enumeration of individuals in functional categories is proposed as a simplified method for studying the abundance and activity of protozoa in ecosystems. Examination of changes in functional group composition and the relationship of functional group abundances to rates of carbon processing are suggested for studies of the importance of protozoa to the flow of energy and materials in ecosystems.     

Allometries of the durations of torpid and euthermic intervals during mammalian hibernation: A test of the theory of metabolic control of the timing of changes in body temperature

Summary The durations of the intervals of torpor and euthermia during mammalian hibernation were found to be dependent on body mass. These relationships support the concept that the timing of body temperature changes is controlled by some metabolic process. Data were obtained from species spanning nearly three orders of magnitude in size, that were able to hibernate for over six months without food at 5°C. The timing of body temperature changes was determined from the records of copper-constantan thermocouples placed directly underneath each animal. Because all species underwent seasonal changes in their patterns of hibernation, animals were compared in midwinter when the duration of euthermic intervals was short and relatively constant and when the duration of torpid intervals was at its longest. Large hibernators remained euthermic longer than small hibernators (Fig. 2). This was true among and within species. The duration of euthermic intervals increased with mass at the same rate (mass^0.38) that mass-specific rates of euthermic metabolism decrease, suggesting that hibernators remain at high body temperatures until a fixed amount of metabolism has been completed. These data are consistent with the theory that each interval of euthermia is necessary to restore some metabolic imbalance that developed during the previous bout of torpor. In addition, small species remained torpid for longer intervals, than large species (Fig. 3). The absolute differences between different-sized species were large, but, on a proportional basis, they were comparatively slight. Mass-specific rates of metabolism during torpor also appear to be much less dependent on body mass than those during euthermia, but the precision of these metabolic measurements is insufficient for them to provide a conclusive test of the metabolic theory. Finally, small species with high mass-specific rates of euthermic metabolism are under tighter energetic constraints during dormancy than large species. The data presented here show that, in midwinter, small species compensate both by spending less time at high body temperatures following each arousal episode and by arousing less frequently, although the former is far more important energetically than the latter.     

Energetically costly behaviour and the evolution of resting metabolic rate in insects

1. A general hypothesis is presented to explain interspecific differences in size-independent resting metabolic rate. This hypothesis is based on a presumed trade-off between a low resting metabolism and adaptations of metabolism during activity.
2. With such a trade-off, selection to reduce resting metabolism is less intense in active species than in species where resting metabolism constitutes a large proportion of the daily metabolic costs. Those animals that spend more energy on activity should therefore have a higher resting metabolic rate than animals that spend less energy on activity.
3. A literature review reveals that flying insects have higher resting metabolic rates than species that use energetically less demanding types of locomotion.
4. Insects producing acoustic advertisement signals can be shown to have higher mass-independent resting metabolic rates than closely related species without this energetically demanding behaviour.
5. Literature data on vertebrate resting metabolic rates are also consistent with the presented hypothesis: the more energy animals spend on activity, the higher the mass-independent resting metabolic rate.     

Survival of ciliate protozoa under starvation conditions and at low bacterial levels

Under starvation conditions, 50% survivorship times displayed no significant relationship with cell size in 2 ciliate species in this study and 5 protozoan species from the literature. Differences in survival ability were attributed to differences in weight-specific respiratory rate and relative motility among these 7 species. At low bacterial levels, 4 ciliate species in this study displayed significant differences in survivorship. High survivorship ofEuplotes patella relative to that ofParamecium caudatum andParaurostyla sp. at low ciliate densities was attributed to the lower individual energy requirements of this smaller species. High survivorship ofStentor coeruleus was interpreted as an effect of its large quantity of reserves and low respiratory rate. The survivorship ofE. patella was reduced at a higher population density. Four ciliate species survived longer at 15C than at 22C. Q₁₀ values based on 50% survivorship times at these 2 temperatures were much lower than Q₁₀ values based on respiratory rates and growth rates of well-fed ciliates over a similar temperature range.     

Some physiological aspects of the autecology of the suspension-feeding protozoanTetrahymena pyriformis

Feeding, growth, and reproductive responses of the suspension-feeding protozoanTetrahymena pyriformis to shifts up or down of the density of its bacterial food were observed. The rates of feeding, growth, and reproduction were determined by measuring the rates of uptake of viable bacterial cells, of change of mean volume of the protozoan cells, and of change of number of protozoan cells, respectively. The effects of the nutritional status of the protozoans at the time of shifting were observed also. Results are interpreted in terms of the limited polymorphism exhibited in the life cycle of this organism. Responses in all cases seem to reflect a strategy for exploiting a patchy, transient environment, a conclusion already reached by several earlier investigators.     

Age at first reproduction and growth rate are independent of basal metabolic rate in mammals

This study tested an emergent prediction from the Metabolic Theory of Ecology (MTE) that the age at first reproduction (α) of a mammal is proportional to the inverse of its mass-corrected basal metabolic rate: The hypothesis was tested with multiple regression models of conventional species data and phylogenetically independent contrasts of 121 mammal species. Since age at first reproduction is directly influenced by an individual’s growth rate, the hypothesis that growth rate is proportional to BMR was also tested. Although the overall multiple regression model was significant, age at first reproduction was not partially correlated with either body mass, growth rate or BMR. Similarly, growth rate was not correlated with BMR. Thus at least for mammals in general, there is no evidence to support the fundamental premise of the MTE that individual metabolism governs the rate at which energy is converted to growth and reproduction at the species level. The exponents of the BMR allometry calculated using phylogenetic generalized least squares regression models were significantly lower than the three-quarter value predicted by the MTE. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Energy expenditure and protein turnover in three species of wallabies (Marsupialia: Macropodidae)

Summary Relationships between basal and fed metabolic rates and whole-body protein turnover rates were examined in three species of wallabies, the red-necked pademelon (Thylogale thetis), parma wallaby (Macropus parma) and tammar wallaby (M. eugenii).There were no significant differences among wallaby species in basal metabolic rate (BMR) which was 30% below eutherian mammals. However, the fed metabolic rate of the tammar was lower than that of the other two species (P<0.05), as was the protein turnover rate (P<0.01) which is consistent with its lower voluntary feed intake and with its lower maintenance nitrogen requirement.Protein turnover rates in the wallabies were 23–47% lower than in eutherian mammals. Similarly, protein synthesis made a lower contribution to fed metabolic rates in the wallabies (7–8%) than in eutherians (17–25%).Thus, compared with several eutherian species, macropodid marsupials have low rates of both energy and protein metabolism, but within the macropodids there is not necessarily a close link between basal metabolic rate and whole-body protein turnover.Abbreviations BMR basal metabolic rate - DEE daily energy expenditure - EE energy expenditure - LSD least significant difference - RQ respiratory quotient     

The cost of sex: quantifying energetic investment in gamete production by males and females

The relative energetic investment in reproduction between the sexes forms the basis of sexual selection and life history theories in evolutionary biology. It is often assumed that males invest considerably less in gametes than females, but quantifying the energetic cost of gamete production in both sexes has remained a difficult challenge. For a broad diversity of species (invertebrates, reptiles, amphibians, fishes, birds, and mammals), we compared the cost of gamete production between the sexes in terms of the investment in gonad tissue and the rate of gamete biomass production. Investment in gonad biomass was nearly proportional to body mass in both sexes, but gamete biomass production rate was approximately two to four orders of magnitude higher in females. In both males and females, gamete biomass production rate increased with organism mass as a power law, much like individual metabolic rate. This suggests that whole-organism energetics may act as a primary constraint on gamete production among species. Residual variation in sperm production rate was positively correlated with relative testes size. Together, these results suggest that understanding the heterogeneity in rates of gamete production among species requires joint consideration of the effects of gonad mass and metabolism.     

Metabolic level and size scaling of rates of respiration and growth in unicellular organisms

1.  Metabolic rate is conventionally assumed to scale with body mass to the 3/4-power, independently of the metabolic level of the organisms being considered. However, recent analyses in a variety of animals and plants indicate that the power (log–log slope) of this relationship varies significantly with metabolic level, ranging from c . 2/3 to 1.
2.  Here I show that the scaling slopes of rates of respiration and growth are related to the metabolic level of a variety of unicellular organisms, as similarly occurs for respiration rates in multicellular organisms.
3.  The recently proposed 'metabolic-level boundaries hypothesis' provides insight into these effects of metabolic level. As predicted, the scaling slopes for resting (endogenous) respiration rate in prokaryotes, algae and protozoans are negatively related to metabolic level; and in protozoans, the scaling slope increases with starvation. Also as predicted, the scaling slopes of growth rate in algae and protozoans are negatively related to growth level. Unexpectedly, opposite effects of starvation on the metabolic scaling slopes of unicellular prokaryotes (compared to that of eukaryotes) may be a spurious result of respiration measurements that did not adequately consider the effects of rapid cell multiplication in prokaryotes with extremely short generation times.
4.  Analyses of both unicellular and multicellular organisms show that there is no universal metabolic scaling relationship, and that variation in metabolic scaling relationships is systematically and possibly universally related to metabolic level.     

Metabolic rate does not scale with body mass in cultured mammalian cells

The allometric scaling of metabolic rate with organism body mass can be partially accounted for by differences in cellular metabolic rates. For example, hepatocytes isolated from horses consume almost 10-fold less oxygen per unit time as mouse hepatocytes [Porter and Brand, Am J Physiol Regul Integr Comp Physiol 269: R226-R228, 1995]. This could reflect a genetically programmed, species-specific, intrinsic metabolic rate set point, or simply the adaptation of individual cells to their particular in situ environment (i.e., within the organism). We studied cultured cell lines derived from 10 mammalian species with donor body masses ranging from 5 to 600,000 g to determine whether cells propagated in an identical environment (media) exhibited metabolic rate scaling. Neither metabolic rate nor the maximal activities of key enzymes of oxidative or anaerobic metabolism scaled significantly with donor body mass in cultured cells, indicating the absence of intrinsic, species-specific, cellular metabolic rate set points. Furthermore, we suggest that changes in the metabolic rates of isolated cells probably occur within 24 h and involve a reduction of cellular metabolism toward values observed in lower metabolic rate organisms. The rate of oxygen delivery has been proposed to limit cellular metabolic rates in larger organisms. To examine the effect of oxygen on steady-state cellular respiration rates, we grew cells under a variety of physiologically relevant oxygen regimens. Long-term exposure to higher medium oxygen levels increased respiration rates of all cells, consistent with the hypothesis that higher rates of oxygen delivery in smaller mammals might increase cellular metabolic rates.     

Effects of Furfural on the Respiratory Metabolism of Saccharomyces cerevisiae in Glucose-Limited Chemostats

Effects of furfural on the aerobic metabolism of the yeast Saccharomyces cerevisiae were studied by performing chemostat experiments, and the kinetics of furfural conversion was analyzed by performing dynamic experiments. Furfural, an important inhibitor present in lignocellulosic hydrolysates, was shown to have an inhibitory effect on yeast cells growing respiratively which was much greater than the inhibitory effect previously observed for anaerobically growing yeast cells. The residual furfural concentration in the bioreactor was close to zero at all steady states obtained, and it was found that furfural was exclusively converted to furoic acid during respiratory growth. A metabolic flux analysis showed that furfural affected fluxes involved in energy metabolism. There was a 50% increase in the specific respiratory activity at the highest steady-state furfural conversion rate. Higher furfural conversion rates, obtained during pulse additions of furfural, resulted in respirofermentative metabolism, a decrease in the biomass yield, and formation of furfuryl alcohol in addition to furoic acid. Under anaerobic conditions, reduction of furfural partially replaced glycerol formation as a way to regenerate NAD⁺. At concentrations above the inlet concentration of furfural, which resulted in complete replacement of glycerol formation by furfuryl alcohol production, washout occurred. Similarly, when the maximum rate of oxidative conversion of furfural to furoic acid was exceeded aerobically, washout occurred. Thus, during both aerobic growth and anaerobic growth, the ability to tolerate furfural appears to be directly coupled to the ability to convert furfural to less inhibitory compounds.