SIZE-DEPENDENT PHOSPHORUS UPTAKE KINETICS AND CELL QUOTA IN PHYTOPLANKTON1

The allometric equation, y = aX^b, described the interspecific variation of phosphate uptake kinetics and cell quota with phytoplankton cell size and showed that smaller cells are superior in uptake rate to large. Species-specific measurements, made by track autoradiography in phosphorus deficient cultures of communities from a phosphorus-limited lake, revealed that eight different species did not differ significantly in the Michaelis-Menten half-saturation constant, K_m. However, both saturated uptake rates (V_max) and the initial slope of the uptake curve (V_max:K_m) decreased per unit biomass with increasing cell size. Biomass-specific cell phosphorus quotas also decreased with increasing cell volume, but less rapidly than did V_max or V_max: K_m. Comparable data from the literature showed that marine species were superior in phosphorus uptake to freshwater species of similar size, but allometric variation of kinetics appeared to exist within both groups. Together with a variable internal stores model of phosphorus-limited growth, the allometric relationships of uptake kinetics and quotas predicted competition to favor smaller cells, with a differential in growth rate diminishing as competitive intensity increased.     

Increasing importance of small phytoplankton in a warmer ocean

The macroecological relationships among marine phytoplankton total cell density, community size structure and temperature have lacked a theoretical explanation. The tiniest members of this planktonic group comprise cyanobacteria and eukaryotic algae smaller than 2 μm in diameter, collectively known as picophytoplankton. We combine here two ecological rules, the temperature–size relationship with the allometric size‐scaling of population abundance to explain a remarkably consistent pattern of increasing picophytoplankton biomass with temperature over the ?0.6 to 22 °C range in a merged dataset obtained in the eastern and western temperate North Atlantic Ocean across a diverse range of environmental conditions. Our results show that temperature alone was able to explain 73% of the variance in the relative contribution of small cells to total phytoplankton biomass regardless of differences in trophic status or inorganic nutrient loading. Our analysis predicts a gradual shift toward smaller primary producers in a warmer ocean. Because the fate of photosynthesized organic carbon largely depends on phytoplankton size, we anticipate future alterations in the functioning of oceanic ecosystems.     

INFLUENCE OF IRRADIANCE ON CELL VOLUME AND CARBON QUOTA FOR TEN SPECIES OF MARINE PHYTOPLANKTON1

Ten species of marine phytoplankton were grown under a range of photosynthetic photon flux densities (PFDs) and examined for variation in cell volume and carbon quota. Results suggest that in response to low PFDs phytoplankton generally reduce their cell volume and frequently reduce their carbon quota. A significant linear relationship between the log of PFD (I) and cell volume (in nine of ten species) and log I and carbon quota (four of ten species) was demonstrated. When exposed, to a transient in light intensity, Thalassiosira pseudonana (Hustedt, clone 3H) Hasle and Heimdal underwent a rapid adaptation in cell volume and carbon quota. Cells going from low light to high light reached maximum mean cell volume within 5 h, and cells going from high light to low light reached a minimum mean cell volume within 12 h. The resulting kinetic constant (k; a measure of the rate of adaptation) was considerably larger than previously reported k values. Ditylum brightwellii (West) Grunow increased in length but did not increase in width during a transient to increased irradiance. Nutrient limitation was shown to override PFD in determining cell volume and carbon quota for Heterosigma akashiwo Hada. Cells grown at equivalent irradiances but N-limited, were smaller than light-limited and nutrient-saturated cells. Therefore, cell volume and carbon quota do not have the same relationship with PFD when factors other than PFD control growth rate. The ecological implications of reduced cell volumes and carbon quotas with decreasing PFD include possible impacts on CO₂ budgets, an influence on sinking rates, potential changes in predation rates, and surface area/cell volume benefits.     

A MODEL APPROACH FOR SIZE-SELECTIVE COMPETITION OF MARINE PHYTOPLANKTON FOR FLUCTUATING NITRATE AND AMMONIUM1

Phytoplankton size-selective competition for fluctuating nutrients was studied with the use of a numerical model, which describes nitrate and ammonium uptake, nitrate reduction to ammonium, and growth as a function of cell she under fluctuating nitrogen limitation. The only size-dependent parameter in the model was the cell nutrient quota. Related to this, the cell surface area per biomass was negatively correlated to cell volume, and the vacuole volume per biomass ratio was positively correlated to cell volume. Simulations showed an inverse correlation between the maximum specific growth rate and cell size under steady-state conditions. With nitrate as the limiting nitrogen source, nitrogen quotas were always higher than with ammonium at the same specific growth rate. Net passive transport of ammonium due to unspecific diffusion of ammonia across the plasma membrane decreased the affinity for ammonium, whereas the affinity for nitrate was not influenced. Transient state-specific ammonium uptake was not dependent on cell size. However, small algae always have the highest specific growth rate in ammonium-controlled systems according to our model. Transient state nitrate uptake rate was positively correlated to cell size because larger algae have a higher vacuole volume per biomass, in which nitrate can be stored. Despite their lower maximum growth rate, larger algae became dominant during simulations under fluctuating nitrate supply when amplitude of and the period between nitrate pulses were high enough. Results from model simulations were qualitatively validated by earlier observations that large diatoms become dominant under fluctuating conditions when nitrate is the main nitrogen source.     

NON-STEADY STATE DYNAMICS OF ALGAL POPULATION GROWTH: EXPERIMENTS WITH TWO CHLOROPHYTES1

The relations among dissolved phosphorus, cell quota of phosphorus, and population growth rate were determined for two Chlorophytes, Chlorella sp. and Scenedesmus quadricauda var. longispina (Chod.) G. M. Smith, in two types of non-steady state continuous culture. One of these types had relatively smooth transitions between growth under different degrees of phosphorus limitation. Under these conditions, two equations often applied to growth kinetics in steady state cultures were found to apply to non-steady state growth. Monad's equation described the relation between dissolved phosphorus concentration and population growth rate, and Droop's equation described the relation between cell quota and population growth rate. The second type of culture received phosphorus only as periodic pulses, leading to sharp changes in dissolved phosphorus, cell quota, and growth rate. A simulation model based on Droop's equation described much of the observed dynamics of cell numbers and quotas in these cultures. Droop's equation could not be convincingly fitted directly to the data, however, due to its incorrect prediction of an immediate growth response to phosphorus pulses. A third relation, predicting that saturated rates of phosphorus uptake would depend on the recent nutrient history of the cells as reflected by the cell quota, was not supported.     

阔叶红松林下6种早夏草本不同生长期生物量分配及模型构建

早夏草本植物作为阔叶红松林林下的一类优势物种,对整个生长季林下草本物种多样性和生物量具有重要影响,对其不同生长阶段生物量分配特征及生物量模型的研究有助于了解该类植物生存策略和碳汇储量。以吉林省蛟河地区阔叶红松林林下的白花碎米荠(Cardamine leucantha)、北重楼(Paris verticillata)、鹿药(Smilacina japonica)、美汉草(Meehania fargesii)、山茄子(Anisodus acutangulus)和紫花变豆菜(Sanicula rubriflora)6种早夏草本植物为研究对象,从4月初到8月底对其物候期进行观察记录,定期取样,从而对6种植物不同生长时期各组分生物量分配特征,地上与地下生物量相对生长关系进行分析研究,并以株高级为自变量建立5种形式(一元线性模型、一元二次模型、指数模型、幂函数模型、对数模型)的单种和混种生物量模型,选取最优模型进行验证。结果表明,6种早夏草本植物花期一般开始于4月底结束于6月中旬,果期开始于5月底结束于8月中旬,不同植物的花、果期持续时间差异较大。生长期内,随着植物生长,株高、生物量和根冠比(R/S)变化明显,但变化趋势不一致。不同物种各组分生物量分配存在差异,用于繁殖的生物量分配比例较小,通常不超过5%。所有物种AGB和BGB间均具有明显的相对生长关系(P0.0001),且均表现为异速生长(相关生长指数a≠1)。根据R~2和SEE选取最优生物量模型,其中幂函数模型形式最常用,其次是一元二次和指数模型。所有最优模型的R~2均较高且SEE较低,拟合效果较好,其中AGB和TB的最优模型要优于BGB,单种模型优于混种模型;通过验证,除混合模型BGB的RMA(30.679%)稍大于30%外,所有模型的RS、EE和RMA均小于30%,P值均大于80%,说明所建立的最优模型能够用来对该地区阔叶红松林林下早夏草本植物生物量进行估算。     

武汉东湖的磷-浮游植物动态模型

本文报道了东湖的一个富营养化模型。这个模型按照1年的时间标度描述东湖藻类的生长和磷循环,其状态变量包括浮游植物磷、藻类生物量、正磷酸盐、碎屑磷和沉积物磷。模型校准和检验结果表明,模型对于系统给定状态的描述是令人满意的,并且对于系统的强制函数的改变能给予合理的响应。根据东湖富营养化工程治理的初步设想,利用模型进行了东湖污水截流前后的水质预报,同时考查了截流后移出沉积物或引灌江水对于改善其水质的效果。模型所提供的各种预报可供拟定东湖治理方案时参考。     

An Examination of Cell Volume in Dominant Phytoplankton Species of the Central and Southern Adriatic Sea

The phytoplankton biomass expressed as total cell volume is often of doubtful value due to cell size and volume variations in the organisms present. Cell volume calculations for 95 phytoplankton species and volume-frequency diagrams for 25 dominant species were analyzed for the Adriatic Sea. Recommendations for more precise work dealing with phytoplankton cell volume and biomass are given. Temporal variations in the average microplankton cell volume are presented and discussed.     

Succession of phytoplankton species in indoor reservoir models varying in sediment composition and water inlet site

Using four continuous-flow indoor reservoir models, the combined effects of varying water inlet depth and sediment composition on phytoplankton species composition and its succession pattern were studied. Species which are known to bloom in eutrophicated reservoirs were dominant in the species succession in each tank. At any one time, a step-like pattern was observed in the composition of the dominant species of each tank, wherein all the species which appeared in a tank with a lower phosphorus concentration were included in the species composition of the tank with a higher phosphorus concentration in an additive relationship.     

Brain Size and Number of Neurons: An Exercise in Synthetic Neuroanatomy

Certain remarkable invariances have long been known in comparative neuroanatomy, such as the proportionality between neuronal density and the inverse of the cubic root of brain volume or that between the square root of brain weight and the cubic root of body weight. Very likely these quantitative relations reflect some general principles of the architecture of neuronal networks. Under the assumption that most of brain volume is due to fibers, we propose four abstract models: I, constant fiber length per neuron; II, fiber length proportionate to brain diameter; III, complete set of connections between all neurons; IV, complete set of connections between compartments each containing the square root of the total number of neurons. Model I conforms well to the cerebellar cortex. Model II yields the observed comparative invariances between number of neurons and brain size. Model III is totally unrealistic, while Model IV is compatible with the volume of the hemispheric white substance in different mammalian species.     

Phytoplankton structure and dynamics in Lake Sanabria and Valparaíso reservoir (NW Spain)

The aim of this work is to compare the composition and seasonality of the phytoplankton population in a natural oligotrophic lake (Lake Sanabria) and a mesotrophic reservoir (Valparaíso). Both ecosystems are located on the Tera river course (NW Spain), which runs along an area of ancient metamorphic and plutonic rocks. Some physical and chemical parameters, chlorophyll a and phytoplankton biovolume were studied from monthly samples collected at different depths during the periods 1987–1989 (Lake Sanabria) and 1991–1992 (Valparaíso). Phytoplankton biovolume and chlorophyll a concentration were about five times higher in Valparaíso than in Lake Sanabria. Species composition (and main phytoplankton groups) were different. Valparaíso was highly dominated by diatoms and Lake Sanabria by cryptophytes and small chlorophytes. In spite of the fact that both sites were nitrogen limited, heterocystous cyanophytes (Anabaena sp.) were detected only in Valparaíso. The relationships between phytoplankton structure and trophic level, hydrological conditions and nitrate content are discussed.     

Kolmogorov-type competition model with finitely supported allocation profiles and its applications to plant competition for sunlight

A Kolmogorov-type competition model featuring allocation profiles, gain functions, and cost parameters is examined. For plant species that compete for sunlight according to the canopy partitioning model [R.R. Vance and A.L. Nevai, Plant population growth and competition in a light gradient: a mathematical model of canopy partitioning, J. Theor. Biol. 245 (2007), pp. 210–219] the allocation profiles describe vertical leaf placement, the gain functions represent rates of leaf photosynthesis at different heights, and the cost parameters signify the energetic expense of maintaining tall stems necessary for gaining a competitive advantage in the light gradient. The allocation profiles studied here, being supported on three alternating intervals, determine “interior” and “exterior” species. When the allocation profile of the interior species is a delta function (a big leaf) then either competitive exclusion or coexistence at a single globally attracting equilibrium point occurs. However, if the allocation profile of the interior species is piecewise continuous or a weighted sum of delta functions (multiple big leaves) then multiple coexistence states may also occur.     

FURTHER EVIDENCE FOR ANOMALOUS SIZE SCALING OF RESPIRATION IN PHYTOPLANKTON1

Respiration per unit mass decreases as organism size increases among metazoans and heterotrophic unicells. The rate of decrease is described by a power function of organism mass; the exponent of the power function is 0.75 (Three-fourths Rule). Previously unanalyzed respiration rates for 11 species of phytoplankton ranging in size over four orders of magnitude show a size-scaling exponent of 1.13 (SE, ±0.15), which is statistically different from 0.75. This result confirms the result of an earlier study of eight phytoplankton species indicating that size scaling of respiration is absent or minimal in phytoplankton, in contrast to the pattern of heterotrophic unicells. The size-related range of respiration rates per unit mass across the full size spectrum of phytoplankton would be approximately 18–fold if respiration were scaled according to the Three-fourths Rule. If respiration does not scale with size or scales minimally with size, as suggested by present evidence, the size-related range of rates will be much smaller or negligible. The apparent anomaly of size scaling for phytoplankton respiration is potentially of great ecological and adaptive significance in unicellular algae.