Toxin production in migrating dinoflagellates: a modelling study of PSP producing Alexandrium

A mechanistic model of dinoflagellate physiology, previously developed and parameterised to simulate paralytic shellfish poison (PSP) content and cell growth for Alexandrium fundyense in response to N and P nutrition, was operated within a vertical water structure in which the organism migrated. Simulations showed the expected development of vertical migration behaviour in response to light and mineral nutrient interactions. Growth in a N-limited water column resulted in a continual, though low level, PSP production with a large population biomass. A sequence of P-stress and nutrient re-feeding during vertical migration stimulated an enhancement of PSP content even with only moderately elevated supply of N:P ratios. This was exacerbated by low absolute P concentrations below the nutricline as well as by the N:P ratio. Although the final biomass was lower in these P-limited simulations, the total toxin production was much higher. The simulations suggest that vertical migration in stratified waters in even moderately high N:P waters could result in the formation of highly toxic populations of Alexandrium. One may expect a similar enhancement of toxicity in other harmful algal species that are engaged in vertical migration, where nutrient supply ratios affect toxin production.     

Simulation of carbon reserve dynamics in Microcystis and its influence on vertical migration with Yoyo model

Blue-green algae control their buoyancy depending upon the surrounding conditions. This process is essential for Cyanobacteria development and can account for their dominance in eutrophic waters in summer. In order to determine the main regulating factors of those movements, we developed a mechanistic and deterministic model, based on differential equations, that simulates the vertical migration of Microcystis sp. In Microcystis, buoyancy regulation results from the dynamics of the carbohydrate reserve metabolism during photosynthesis. These fundamental processes are modelled daily by this vertical 1-D model named Yoyo. It describes the movement of colonies with different sizes in response to variations of environmental conditions. This paper presents the model sensitivity analysis. We individually investigated the role of light and temperature upon algal migration with colonies of two different diameters. Under a daily light cycle and a temperature of 20 degrees C, the model described vertical migration on a 48 h rhythm in colonies with a 300-micron diameter.     

Computer simulations of seasonal outbreak and diurnal vertical migration of cyanobacteria

Algal blooms caused by cyanobacteria are characterized by two features with different time scales: one is seasonal outbreak and collapse of a bloom and the other is diurnal vertical migration. Our two-component mathematical model can simulate both phenomena, in which the state variables are nutrients and cyanobacteria. The model is a set of one-dimensional reaction-advection-diffusion equations, and temporal changes of these two variables are regulated by the following five factors: (1) annual variation of light intensity, (2) diurnal variation of light intensity, (3) annual variation of water temperature, (4) thermal stratification within a water column and (5) the buoyancy regulation mechanism. The seasonal change of cyanobacteria biomass is mainly controlled by factors, (1), (3) and (4), among which annual variations of light intensity and water temperature directly affect the maximum growth rate of cyanobacteria. The latter also contributes to formation of the thermocline during the summer season. Thermal stratification causes a reduction in vertical diffusion and largely prevents mixing of both nutrients and cyanobacteria between the epilimnion and the hypolimnion. Meanwhile, the other two factors, (2) and (5), play a significant role in diurnal vertical migration of cyanobacteria. A key mechanism of vertical migration is buoyancy regulation due to gas-vesicle synthesis and ballast formation, by which a quick reversal between floating and sinking becomes possible within a water column. The mechanism of bloom formation controlled by these five factors is integrated into the one-dimensional model consisting of two reaction-advection-diffusion equations.     

OBSERVATIONS ON THE SWIMMING AND BUOYANCY OF SOME THECOSOMATOUS PTEROPOD GASTROPODS

Cymbulia peroni produces forward propulsion only on the upstrokeof its ‘wingbeat’ the downstroke opposing slightnegative buoyancy. The species' torpedo-shaped pseudoconchaprovides stability as well as buoyancy. Corolla ovata producespropulsive power on the downstroke of its ’wings‘and shows periods of active upward swimming alternating withpassive downward gliding/parachuting. Both cymbuliid speciesare gelatinous slow movers, close to neutral buoyancy, and areincapable of effective diurnal vertical migration. Two cavoliniidspecies were also studied; Cilo pyramida and Cuvierina columnella.Both were dense and sank quickly if they ceased active swimming.Both had postural responses to slow sinking, but vigorous activitywas required to hover in the water column. Clio could not climbrapidly enough to carry out effective diurnal vertical migration,but was capable of fast horizontal swimming during which thetriangular shell appeared to act as a hydrofoil. The smallerCuvierina could not swim quickly in a horizontal direction,but was capable of swimming vertically at rates sufficient tomaintain station with moving isolumes and may therefore carryout significant diurnal vertical migrations. (Received 28 September 1989; accepted 12 December 1989)     

Stay dormant or escape sprouting? Turion buoyancy and sprouting abilities of the submerged macrophyte Potamogeton crispus L.

The ability of asexual propagules to disperse is an important ecological determinant of the spread and establishment of many aquatic species. However, few previous studies have addressed the relationship between the asexual propagule buoyancy and sprouting abilities in submerged macrophytes. For this reason, turions of Potamogeton crispus samples were collected from Lake Liangzi, and an incubator sprouting experiment was conducted. Our results revealed that the floating turions showed higher sprouting rates than that of sinking turions, indicating the former ones are possibly with high levels of primary metabolites. The higher N and P concentrations in the floating turions caused lower C:N, C:P, and N:P ratios in these turions compared with sinking turions, which confirmed the activation of floating turions. The free amino acid and soluble carbohydrate concentrations were also higher in floating turions than those in sinking turions. Our results also revealed that turion leaf porosity rather than starch concentration may determine the density of P. crispus turions. This study makes a contribution to our understanding of how the internal characteristics of turions can (at least partly) determine dispersal outcomes and offers new insights into the dispersal and sprouting of asexual propagules of submerged macrophytes.     

The development and vertical distribution of populations of gas-vacuolate bacteria in a eutrophic,monomictic lake

The role of gas vacuoles in the vertical stratification of planktonic bacteria is analysed. Measurements made with certain gas-vacuolate bacteria in laboratory culture suggest that only colonial forms could sink or float fast enough to form population maxima in lakes by vertical migration from other depths. It is suggested that in the case of individual cells the importance of the buoyancy provided by gas vacuoles is to minimise sinking rates and thereby to increase residence times of the organisms at depths where conditions support their growth.Changes in the vertical distribution of a number of gas-vacuolate bacteria were followed throughout the year in a monomictic, eutrophic lake (Crose Mere, Shropshire). All were restricted to the anaerobic hypolimnion which developed in summer. The various species formed maxima at different depths and times. With some of them (e.g. species of Thiopedia, Pelonema and Brachyarcus) growth was necessary to explain their development. In others (e. g. species of Pelodictyon and two colourless bacteria) vertical migrations might also have contributed to their development.     

Redox control of N:P ratios in aquatic ecosystems

The ratio of dissolved fixed inorganic nitrogen to soluble inorganic phosphate (N:P) in the ocean interior is relatively constant, averaging ~16 : 1 by atoms. In contrast, the ratio of these two elements spans more than six orders of magnitude in lakes and other aquatic environments. To understand the factors influencing N:P ratios in aquatic environments, we analyzed 111 observational datasets derived from 35 water bodies, ranging from small lakes to ocean basins. Our results reveal that N:P ratios are highly correlated with the concentration of dissolved O₂ below ~100 µmol L⁻¹. At higher concentrations of O₂, N:P ratios are highly variable and not correlated with O₂; however, the coefficient of variation in N:P ratios is strongly related to the size of the water body. Hence, classical Redfield ratios observed in the ocean are anomalous; this specific elemental stoichiometry emerges not only as a consequence of the elemental ratio of the sinking flux of organic matter, but also as a result of the size of the basins and their ventilation. We propose that the link between N:P ratios, basin size and oxygen levels, along with the previously determined relationship between sedimentary δ¹⁵N and oxygen, can be used to infer historical N:P ratios for any water body.     

Unexpected Positive Buoyancy in Deep Sea Sharks,Hexanchus griseus,and a Echinorhinus cookei

We do not expect non air-breathing aquatic animals to exhibit positive buoyancy. Sharks, for example, rely on oil-filled livers instead of gas-filled swim bladders to increase their buoyancy, but are nonetheless ubiquitously regarded as either negatively or neutrally buoyant. Deep-sea sharks have particularly large, oil-filled livers, and are believed to be neutrally buoyant in their natural habitat, but this has never been confirmed. To empirically determine the buoyancy status of two species of deep-sea sharks (bluntnose sixgill sharks, Hexanchus griseus, and a prickly shark, Echinorhinus cookei) in their natural habitat, we used accelerometer-magnetometer data loggers to measure their swimming performance. Both species of deep-sea sharks showed similar diel vertical migrations: they swam at depths of 200–300 m at night and deeper than 500 m during the day. Ambient water temperature was around 15°C at 200–300 m but below 7°C at depths greater than 500 m. During vertical movements, all deep-sea sharks showed higher swimming efforts during descent than ascent to maintain a given swimming speed, and were able to glide uphill for extended periods (several minutes), indicating that these deep-sea sharks are in fact positively buoyant in their natural habitats. This positive buoyancy may adaptive for stealthy hunting (i.e. upward gliding to surprise prey from underneath) or may facilitate evening upward migrations when muscle temperatures are coolest, and swimming most sluggish, after spending the day in deep, cold water. Positive buoyancy could potentially be widespread in fish conducting daily vertical migration in deep-sea habitats.     

Energy cost and putative benefits of cellular mechanisms modulating buoyancy in aflagellate marine phytoplankton

Little information is available on the energetics of buoyancy modulation in aflagellate phytoplankton, which comprises the majority of autotrophic cells found in the ocean. Here, we computed for three aflagellate species of marine phytoplankton (Emiliania huxleyi, Thalassiosira pseudonana, and Ethmodiscus rex) the theoretical minimum energy cost as photons absorbed and nitrogen resource required of the key physiological mechanisms (i.e., replacement of quaternary ammonium by dimethyl‐sulfoniopropionate, storage of polysaccharides, and cell wall biosynthesis) affecting the cell's vertical movement as a function of nitrogen (N) availability. These energy costs were also normalized to the capacity of each buoyancy mechanism to modulate sinking or rising rates based on Stokes' law. The three physiological mechanisms could act as ballast in the three species tested in conditions of low N availability at a low fraction (<12%) of the total photon energy cost for growth. Cell wall formation in E. huxleyi was the least costly ballast strategy, whereas in T. pseudonana, the photon energy cost of the three ballast strategies was similar. In E. rex, carbohydrate storage and mobilization appear to be energetically cheaper than modulations in organic solute synthesis to achieve vertical migration. This supports the carbohydrate‐ballast strategy for vertical migration for this species, but argues against the theory of replacement of low‐ or high‐density organic solutes. This study brings new insights into the energy cost and potential selective advantages of several strategies modulating the buoyancy of aflagellate marine phytoplankton.     

Responses of wetland graminoids to the relative supply of nitrogen and phosphorus

The biomass production of wetland vegetation can be limited by nitrogen or phosphorus. Some species are most abundant in N-limited vegetation, and others in P-limited vegetation, possibly because growth-related traits of these species respond differently to N versus P supply. Two growth experiments were carried out to examine how various morphological and physiological traits respond to the relative supply of N and P, and whether species from sites with contrasting nutrient availability respond differently. In experiment 1, four Carex species were grown in nutrient solutions at five N:P supply ratios (1.7, 5, 15, 45, 135) combined with two levels of supply (geometric means of N and P supply). In experiment 2, two Carex and two grass species were grown in sand at the same .ve N:P supply ratios combined with three levels of supply and two light intensities (45% or 5% daylight). After 12-13 weeks of growth, plant biomass, allocation, leaf area, tissue nutrient concentrations and rates and nutrient uptake depended signi.cantly on the N:P supply ratio, but the type and strength of the responses differed among these traits. The P concentration and the N:P ratio of shoots and roots as well as the rates of N and P uptake were mainly determined by the N:P supply ratio; they showed little or no dependence on the supply level and relatively small interspeci.c variation. By contrast, the N concentration, root mass ratio, leaf dry matter content and speci.c leaf area were only weakly related to the N:P supply ratio; they mainly depended on plant species and light, and partly on overall nutrient supply. Plant biomass was determined by all factors together. Within a level of light and nutrient supply, biomass was generally maximal (i.e. co-limited by N and P) at a N:P supply ratio of 15 or 45. All species responded in a similar way to the N:P supply ratio. In particular, the grass species Phalaris arundinacea and Molinia caerulea showed no differences in response that could clearly explain why P. arundinacea tends to invade P-rich (N-limited) sites, and M. caerulea P-limited sites. This may be due to the short duration of the experiments, which investigated growth and nutrient acquisition but not nutrient con­servation.     

INTERACTING EFFECTS OF LIGHT AND NUTRIENT LIMITATION ON THE GROWTH RATE OF SYNECHOCOCCUS LINEARIS (CYANOPHYCEAE)1

The blue-green alga Synechococcus linearis (Naeg.) Kom. was grown in P- and N-limited chemostats over a range of potentially limiting irradiances in order to determine the combined effects of light and nutrient limitation on some aspects of the composition and metabolism of this alga. Over a narrow range of low irradiances, simultaneous limitation of growth rate by light and either N or P was shown. This simultaneous limitation of growth rate by a nutrient and a physical factor can be explained by the ability of an increased supply of one to compensate in part for a decreased supply of the other. At all irradiances, the internal concentration of the limiting nutrient increased with increasing dilution rate, and the results could be fitted to the Droop relationship. With decreasing irradiance, the internal concentration of the limiting nutrient increased. There appeared to be little or no effect of light on the minimum internal concentration of P but that of N increased with decreasing light. Both chlorophyll a and biliprotein per unit particulate C increased with increasing dilution rate and decreasing irradiance. The critical N/P ratio increased with decreasing light as the N requirement of N-limited cells increased faster than did the P requirement of P-limited cells. The composition of exponentially growing cells in complete medium varied much less with light. Neither dilution rate nor irradiance during growth had a great effect on saturated rates of P or N uptake or alkaline phosphatase activity. Calculated assimilation ratios increased with light and dilution rate. The role of the flexibility of nutrient composition in adaptation to adverse conditions and the implications of the results for the use of physiological indicators of nutrient status are discussed.     

Diel variation of the cellular carbon to nitrogen ratio of Chlorella autotrophica (Chlorophyta) growing in phosphorus‐ and nitrogen‐limited continuous cultures

We investigated the relationship between daily growth rates and diel variation of carbon (C) metabolism and C to nitrogen (N) ratio under P‐ and N‐limitation in the green algae Chlorella autotrophica. To do this, continuous cultures of C. autotrophica were maintained in a cyclostat culture system under 14:10 light:dark cycle over a series of P‐ and N‐limited growth rates. Cell abundance, together with cell size, as reflected by side scatter signal from flow cytometric analysis demonstrated a synchronized diel pattern with cell division occurring at night. Under either type of nutrient limitation, the cellular C:N ratio increased through the light period and decreased through the dark period over all growth rates, indicating a higher diel variation of C metabolism than that of N. Daily average cellular C:N ratios were higher at lower dilution rates under both types of nutrient limitation but cell enlargement was only observed at lower dilution rates under P‐limitation. Carbon specific growth rates during the dark period positively correlated with cellular daily growth rates (dilution rates), with net loss of C during night at the lowest growth rates under N‐limitation. Under P‐limitation, dark C specific growth rates were close to zero at low dilution rates but also exhibited an increasing trend at high dilution rates. In general, diel variations of cellular C:N were low when dark C specific growth rates were high. This result indicated that the fast growing cells performed dark C assimilation at high rates, hence diminished the uncoupling of C and N metabolism at night.     

The Wnt Frizzled Receptor MOM-5 Regulates the UNC-5 Netrin Receptor through Small GTPase-Dependent Signaling to Determine the Polarity of Migrating Cells

Wnt and Netrin signaling regulate diverse essential functions. Using a genetic approach combined with temporal gene expression analysis, we found a regulatory link between the Wnt receptor MOM-5/Frizzled and the UNC-6/Netrin receptor UNC-5. These two receptors play key roles in guiding cell and axon migrations, including the migration of the C. elegans Distal Tip Cells (DTCs). DTCs migrate post-embryonically in three sequential phases: in the first phase along the Antero-Posterior (A/P) axis, in the second, along the Dorso-Ventral (D/V) axis, and in the third, along the A/P axis. Loss of MOM-5/Frizzled function causes third phase A/P polarity reversals of the migrating DTCs. We show that an over-expression of UNC-5 causes similar DTC A/P polarity reversals and that unc-5 deficits markedly suppress the A/P polarity reversals caused by mutations in mom-5/frizzled. This implicates MOM-5/Frizzled as a negative regulator of unc-5. We provide further evidence that small GTPases mediate MOM-5’s regulation of unc-5 such that one outcome of impaired function of small GTPases like CED-10/Rac and MIG-2/RhoG is an increase in unc-5 function. The work presented here demonstrates the existence of cross talk between components of the Netrin and Wnt signaling pathways and provides further insights into the way guidance signaling mechanisms are integrated to orchestrate directed cell migration.     

OPTIMUM N:P RATIOS AND COEXISTENCE OF PLANKTONIC ALGAE1

The optimum atomic ratio of N to P, the ratio at which one nutrient limitation changes over to the other, was determined in seven species of freshwater planktonic algae. The ratio varied over a wide range among species; the average for these species was 17. If the cellular nutrient ratios in marine species are comparable with those in freshwater organisms, Redfield's ratio of 15 is remarkably close to the average. Cellular N:P ratios varied over a 24-h period under a light:dark cycle. The variation of the optimum ratio between species and diel change in cellular N:P ratios within a species could play an important role in population dynamics by enhancing the probability of coexistence of species.     

Interactive effects of N:P ratios and light on nitrogen‐fixer abundance

Nitrogen‐fixers can contribute significant amounts of nitrogen (N) and impact ecosystem functioning in diverse aquatic and terrestrial ecosystems. What determines N‐fixer abundance still remains poorly understood. Here we experimentally investigate major environmental controls on the abundance of N‐fixers: nitrogen to phosphorus (N:P) ratio and light. We grew a N‐fixer, cyanobacterium Anabaena flos‐aquae, in a multispecies community of freshwater phytoplankton in replicated factorial design treatments with two N:P ratios and two light levels. We show that low N:P ratios promote the dominance of the N‐fixer in the community, but only under high light. Under low light, N:P ratio did not have a significant effect on the abundance of the N‐fixer. N fixation occurred at low N:P only and increased with increasing light. In contrast, the density of non N‐fixing cyanobacteria did not depend on N:P ratios. Green algae dominated under high N:P and high light only, exhibiting the opposite pattern of dominance to N‐fixers. These results are consistent with patterns observed in nature and help explain the N‐fixer distribution along the environmental gradients of nutrients and light.     

华北石质山地侧柏人工林C、N、P生态化学计量特征的季节变化

以北京九龙山自然保护区幼龄侧柏人工林为研究对象,对其不同生长季节叶、枝、根(0—10 cm、10—20 cm土层)的碳(C)、氮(N)、磷(P)含量及其生态化学计量学特征进行了分析,深入探讨了生长季节与器官以及两因素交互作用对以上特征的影响,研究有助于理解植物各性状之间的相互作用以及植物生长过程中资源的利用和分配状况。结果表明:1)不同器官间C含量为414.97—461.58 g/kg,枝最大,根(0—10 cm)最小;N含量为6.57—14.28 g/kg,叶最大,枝最小;P含量为0.39—1.28 g/kg,叶最大,根(10—20 cm)最小;C∶N为31.76—70.98,枝最大,叶最小;C∶P为369.93—1099.20,根(10—20 cm)最大,叶最小;N∶P为9.21—23.81,根(0—10 cm)最大,枝最小。整个生长季节中侧柏各器官C含量最稳定,变异系数均小于7%;P含量变异性最大,变异系数均超过15%,N含量变异性介于两者之间;各器官中C∶N和N∶P较C∶P更为稳定,C、N与P具有较好的耦合协同性,C∶P和N∶P的变化主要取决于P的变化。2)器官对C、N、P含量及其化学计量关系均存在显著影响,生长季节对N和P含量存在显著影响,两者交互作用只对P含量存在显著影响,器官对侧柏C、N、P含量及其化学计量变异的贡献大于生长季节。3)侧柏各器官间C、N、P含量及其化学计量比相关性多数未达到显著性水平,仅有叶与枝中的P及C∶P显著相关,说明侧柏器官分化过程中各器官对元素的吸收利用具有特异性。侧柏叶片N∶P14,说明生长季节里幼龄侧柏人工林更多受到N限制。     

ELEMENTAL AND BIOCHEMICAL COMPOSITION OF RHINOMONAS RETICULATA (CRYPTOPHYTA) IN RELATION TO LIGHT AND NITRATE‐TO‐PHOSPHATE SUPPLY RATIOS1

The biochemical basis for variations in the critical nitrogen‐to‐phosphorus (N:P) ratio, which defines the transition between N‐ and P‐limitation of growth rate, is currently not well understood. To assess this issue, we cultured the cryptophyte Rhinomonas reticulata NOVARINO in chemostats with inflow nitrate‐to‐phosphate ratios ranging from 5 to 60 mol N·(mol P)^?1 at two light intensities. The nitrate‐to‐phosphate ratio marking the transition between N‐ and P‐limitation was independent of light intensity and was between 30 and 45 mol N/mol P. In N‐limited cells, the particulate N:P ratio was stable at around 23 mol N/mol P over a range of inflow nitrate‐to‐phosphate from 5 to 30, whereas in P‐limited cells this ratio was around 90 mol N/mol P at inflow nitrate‐to‐phosphate ratios of 45 and 60. Cell phosphorus decreased with increasing nitrate‐to‐phosphate ratio up to the critical nitrate‐to‐phosphate ratio for each light intensity, above which they remained stable. The C:P of R. reticulata cells increased with increasing inflow nitrate‐to‐phosphate from around the Redfield value (106 mol C/mol P) to around 700. There was a significant effect of light on C:P in the N‐ limited cells, with higher C:P under high light conditions that was not observed in the P‐limited chemostats. Cellular RNA was not influenced by light but was greatly influenced by the type of nutrient limitation. In contrast, chl a, C, N, and protein were not influenced by the nitrate‐to‐phosphate in the inflow medium. Total protein per RNA was independent of light intensity but exhibited a maximum at inflow nitrate‐to‐phosphate of 30. Our results suggest a strong “two‐level” homeostatic mechanism of cellular N and P content in R. reticulata with two distinct states that are determined by the type of nutrient limitation and not by light.     

Light, nutrients and primary productivity in Lake Biwa: An evaluation of the current ecosystem situation

Simple correlation and multiple regression analyses were performed to examine the relationship between primary productivity and environmental factors in the north basin of Lake Biwa. The primary production rates used in the analyses were estimated monthly or bimonthly during the growing season (April–November) in 1992, 1996 and 1997 with the ¹³C method. Elemental (C, N and P) contents of seston were used to assess nutrient conditions. Analyses revealed that 86% of variance in depth-integrated primary production rates (areal PP) can be explained by changes in light intensity, and sestonic C, N and P concentrations. Water temperature had no effect on areal PP. To assess relative effects of light and nutrients on PP, the P:B ratio was estimated by normalizing PP with sestonic C. The areal P:B ratio correlated most significantly with the sestonic N:P ratio, followed by light intensity. When regression analyses were made at each depth, however, the P:B ratio correlated significantly only with the sestonic N:P ratio at 0 and 1 m depths, while light intensity was also incorporated into the regressions at deeper than 2.5 m. In these regressions, the P:B ratio was negatively correlated with sestonic N:P ratio but positively with light intensity. The results suggest that the primary production rate in this lake was mainly limited by P relative to N supply rates, but was not free from light limitation in a large part of the epilimnion. In Lake Biwa, the vertical water mixing regime as well as the nutrient supply seem to be important in determining the growth and composition of primary producers, since the surface mixing layer extends into 10–15 m depths during most of the growing season.