Influence of phosphorus concentration on the growth kinetics and stoichiometry of the microalga Scenedesmus obliquus

The growth of the freshwater microalga Scenedesmus obliquus was studied at 30°C in a mineral culture medium with phosphorus concentrations of between 0 and 372 μ . The values for the specific growth rates, between and , fitted a semistructured substrate-limitation model with μ_m1 = 0·0466 h⁻¹, μ_m2 = 0·0256 h⁻¹ and . The specific uptake rate of phosphorus reached a maximum value of q_Sm1 = 658·01 × 10⁻⁴ μmol P mg⁻¹ biomass h⁻¹.     

Plant–herbivore interactions and ecological stoichiometry: when do herbivores determine plant nutrient limitation?

Recent studies on plant–herbivore indirect interactions via nutrient recycling have led to the hypothesis that herbivores with a low nitrogen: phosphorus ratio, feeding on plants with a higher nitrogen: phosphorus ratio, recycle relatively more nitrogen, driving plants into phosphorus limitation. We demonstrate in this paper that such a hypothesis is valid only under restricted conditions, i.e. the nitrogen: phosphorus ratio of inorganic nutrients supplied to the system must be neither too high nor too low compared with the nitrogen: phosphorus ratio of the whole plant + herbivore biomass. If plants have a greater affinity for phosphorus than for nitrogen, low herbivore nitrogen: phosphorus ratio can even promote nitrogen limitation. These results are qualitatively robust, whether grazing functions are donor-controlled or recipient-controlled. We present a graphical analysis of these conditions based on the Zero Net Growth Isocline method.     

Influence of nutrient availability on phytoplankton growth and community structure in the Port Adelaide River,Australia: [2pt] bioassay assessment of potential nutrient limitation

We investigated the influence of nutrient availability, specifically nitrogen, phosphorus and silicon on growth and community structure of phytoplankton from the Port Adelaide River estuary, South Australia. Two bioassay experiments were conducted. The first, Nutrich1, involved addition of nutrients in vitro to samples of the natural phytoplankton community from a single location in the upper estuary. The second, Nutrich2, involved nutrient addition and incubation of water from five locations in the estuary following inoculation with a `standardised' phytoplankton assemblage derived from laboratory cultures. In Nutrich1, enrichment with silicon led to greatly enhanced phytoplankton biomass due to increased growth of diatoms. Addition of nitrogen or phosphorus had little effect on phytoplankton growth. In Nutrich2, addition of nitrogen resulted in enhanced growth of phytoplankton in water collected from near the mouth the estuary, but there were no differences in growth among nutrient treatments for the remaining locations. Comparison of phytoplankton growth rate among locations revealed a trend of decreasing growth in moving towards the mouth of the estuary. This trend was unaffected by enrichment with nitrate, phosphate or silicate. We suggest that spatial variation in growth potential within the Port Adelaide River estuary may relate to variation in the concentration of nitrogen as ammonium.     

Effects of food quality on life history of the rotifer Brachionus calyciflorus Pallas

1. Herbivorous zooplankton face considerable temporal and spatial variation in food quality, to which they respond by adapting their life histories. Zooplankton may even take up mineral nutrients directly, and use these to counter the effects of algal nutrient limitation (mineral compensation). This study examined the life history of the rotifer Brachionus calyciflorus fed phosphorus‐, and nitrogen‐limited Scenedesmus obliquus (Chlorophyta), and investigated whether B. calyciflorus was capable of mineral compensation. 2. Both phosphorus‐ and nitrogen‐limited algae gave similar life history responses: somatic growth and reproduction were reduced, whereas lifespan remained unaffected. 3. No evidence was found for mineral compensation in B. calyciflorus in relation to detrimental life history effects, so mineral compensation does not seem to be relevant for this species under field conditions. 4. The similarity in life history responses of B. calyciflorus and the low levels of ω‐3 PUFAs in both phosphorus‐ and nitrogen‐depleted algae suggest that ω‐3 PUFAs were limiting to B. calyciflorus, although other (bio)chemicals or mineral nutrients may also have been important. 5. No trade‐off was observed between life span and reproduction during algal nutrient limitation. Reduced population growth rates of B. calyciflorus were caused by shorter reproductive periods.     

Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.     

Trophic state and nutrient limitation in Lake Baringo,Kenya

The trophic state of Lake Baringo and factors that could be limiting the development of algal biomass in it were investigated during one wet/dry hydrological cycle in 2014–2015. Water samples were analysed for dissolved inorganic nutrients, including , and , total phosphorus and Chlorophyll a. Light attenuation was estimated using Secchi depth. The trophic state was determined using Carlson trophic state indices (CTSI). Deviations in CTSI, nutrient ratios and ambient nutrient concentrations were used to identify factors limiting phytoplankton growth. The mean values measured for Secchi depth, nitrate, total phosphorus and Chlorophyll a showed significant seasonal variation (p < 0.05). Based on the Carlson trophic state index, the results show that Lake Baringo is eutrophic. However, the lake is also experiencing phosphorus limitation and poor light penetration, because of high turbidity, which is more pronounced during the wet season.     

铜绿微囊藻和斜生栅藻非稳态营养盐限制条件下的生长竞争特性

采用“脉冲”添加方法进行了非稳态条件下铜绿微囊藻（M.）和斜生栅藻（S.）分别在氮磷单营养盐和双营养盐限制时的共培养试验。试验结果显示：当添加频率为1d时,无论何种营养盐限制,M.均成为优势藻种。氮限制条件下,氮时均浓度范围在0.3—2.4 mg/L时,M.始终具有竞争优势。磷限制条件下,磷浓度范围在0.018—0.035 mg/L时,S.只在生长初期阶段占优。氮磷双营养盐限制条件下,添加液的氮磷质量比为35:1（设定为最优比）,添加频率为8 d时,两种藻表现出共生特征;而偏离最优比时（N:P=70:1,17:1）,在不同的添加频率下均未出现共生现象,且氮的时均浓度为0.6—4.8 mg/L时（70:1）,M.具有竞争优势,而降低为0.15—0.3 mg/L时（17:1）,S.占优。随着添加频率的变化,两种藻的细胞大小也会随之改变,S.随着营养盐浓度的降低而增大,且在双营养盐限制条件下变化更显著。上述试验结果分析表明：两种藻竞争能力与添加频率相关,在藻种浓度的变化上,按照‘拾遗－机会’交替竞争理论,M.表现出机会主义者特征,而S.则表现出拾遗者的特征,两者的共生特征也符合‘中度干扰’假说。藻细胞大小变化表明,两种藻均可以改变大小实现最大限度争夺受限营养盐。在低浓度时,S.细胞大小的变化同样也变现出了“拾遗者”的特征。     

Changes in stoichiometric constraints on epilithon and benthic macroinvertebrates in response to slight nutrient enrichment of mountain rivers

1. To assess changes in stoichiometric constraints on stream benthos, we measured elemental composition of epilithon and benthic macroinvertebrates in intrinsically P‐limited mountain rivers, upstream and downstream of low‐level anthropogenic nutrient enrichment by effluents of municipal wastewater treatment plants. 2. While there was a broad range in the elemental composition of epilithon (C : P ratios of 200–16 500, C : N ratios of 8–280, N : P ratios of 8–535) and heptageniid mayfly scrapers (C : P ratios of 125–300, C : N ratios of 5.1–7.2, N : P ratios of 20–60), the average C : P ratio of epilithon was 10‐fold lower and the average C : N ratio twofold lower at more nutrient‐rich downstream sites. Nutrient ratios in benthic macroinvertebrates were lower than in epilithon and varied little between relatively nutrient‐poor and nutrient‐rich sites. 3. We modified the existing definition of producer‐consumer elemental imbalance to allow for variation in consumer nutrient content. We defined this ‘non‐homeostatic’ imbalance as the perpendicular distance between the producer and consumer C : P, C : N, or N : P ratios, and the 1 : 1 line. 4. At P‐limited sites, the estimated mayfly N : P recycling ratio was higher than the N : P ratio in epilithon, suggesting nutrient recycling by consumers could accentuate P‐limitation of epilithon. 5. Measuring the degree of producer–consumer nutrient imbalance may be important in predicting the magnitude of effects from nutrient enrichment and can help elucidate the causes and consequences of ecological patterns and processes in rivers.     

Stoichiometric and growth responses of a freshwater filamentous green alga to varying nutrient supplies: slow and steady wins the race

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Root allocation and multiple nutrient limitation in the New Jersey Pinelands

Abstract Current understanding of the effects of resource stress on plant communities emphasizes the adaptive integration of multiple limiting factors, but it has been difficult to directly demonstrate the fundamental assumption of the adaptive control of limitation. One model predicts a positive correlation between optimal allocation to the uptake of each resource and growth response to enrichment of that resource. Here we report a test of this prediction by a fertilization experiment in the New Jersey Pinelands Biosphere Reserve where limitation was measured by growth response to nutrient enrichments, and allocation was measured by root proliferation in microsites enriched by those same nutrients. Results suggest that a stand of regenerating Pinus echinata Mill. (shortleaf pine) was jointly limited by nitrogen and potassium, with a possible small effect of phosphorus. Root allocation was proportional to growth response, which supports the assumption of adaptive control of limitation.     

太平洋中西部海域浮游植物营养盐的潜在限制

2009年8月至9月期间在太平洋西部N1站和中部N2站进行现场营养盐加富培养实验。结果显示:N1站,浮游植物生物量对N或者P添加都有较强的响应,其中N+P+Si组和N+P组浮游植物长势迅速,叶绿素a从初始的0.03μg/L分别达到2.12μg/L和1.83μg/L,同时P先于N和Si之前被耗尽;说明N1站为N、P共同限制,P是首要限制因子。而N2站,浮游植物生物量仅对N、P共同添加有明显响应,N先于P和Si被浮游植物消耗殆尽。利用培养过程中营养盐比值变化推断,N1站浮游植物以低于Redfield ratio(16N∶1P)吸收N和P;而N2站浮游植物以高于Redfield ratio(16N∶1P)吸收N和P。这可能解释了太平洋西部的寡营养盐海域为潜在P限制,而在太平洋中部海域则为潜在N限制。     

Stoichiometric regulation of phytoplankton toxins

Ecological Stoichiometry theory predicts that the production, elemental structure and cellular content of biomolecules should depend on the relative availability of resources and the elemental composition of their producer organism. We review the extent to which carbon‐ and nitrogen‐rich phytoplankton toxins are regulated by nutrient limitation and cellular stoichiometry. Consistent with theory, we show that nitrogen limitation causes a reduction in the cellular quota of nitrogen‐rich toxins, while phosphorus limitation causes an increase in the most nitrogen‐rich paralytic shellfish poisoning toxin. In addition, we show that the cellular content of nitrogen‐rich toxins increases with increasing cellular N : P ratios. Also consistent with theory, limitation by either nitrogen or phosphorus promotes the C‐rich toxin cell quota or toxicity of phytoplankton cells. These observed relationships may assist in predicting and managing toxin‐producing phytoplankton blooms. Such a stoichiometric regulation of toxins is likely not restricted to phytoplankton, and may well apply to carbon‐ and nitrogen‐rich secondary metabolites produced by bacteria, fungi and plants.     

Contrasting effects of rising CO2 on primary production and ecological stoichiometry at different nutrient levels

Although rising CO₂ concentrations are thought to promote the growth and alter the carbon : nutrient stoichiometry of primary producers, several studies have reported conflicting results. To reconcile these contrasting results, we tested the following hypotheses: rising CO₂ levels (1) will increase phytoplankton biomass more at high nutrient loads than at low nutrient loads, but (2) will increase their carbon : nutrient stoichiometry more at low than at high nutrient loads. We formulated a mathematical model to predict dynamic changes in phytoplankton population density, elemental stoichiometry and inorganic carbon chemistry in response to rising CO₂. The model was tested in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa. The model predictions and experimental results confirmed the hypotheses. Our findings provide a novel theoretical framework to understand and predict effects of rising CO₂ concentrations on primary producers and their nutritional quality as food for herbivores under different nutrient conditions.     

Biotests with phytoplankton assemblages. Growth limitation along temporal and spatial gradients

The development of eight different species (populations) along temporal and vertical gradients in several lakes was studied. Many populations had an exponential growth phase and a decline phase. The growth rate was often high during the exponential phase. Some species, e.g. Oscillatoria spp. and Synedra cf. acus, often also had a long stationary phase. The growth rate and the sinking rate of these populations were often very low. Laboratory batch experiments with dilute phytoplankton populations were carried out to estimate the degree of growth limitation (L) for different populations sample from different lakes during the three growth phases. L was always low and often zero for populations initially in the exponential phase and always high for populations initially in the decline phase. The biotests also gave results that can help to explain the vertical distribution of Oscillatoria or Asterionella in three lakes investigated. The results indicate that the growth rates and the development of the populations were dependent on the external chemical and physical conditions. The transition between the different growth phases seemed often to be dependent on the external nutrient conditions. P, N, Si and Fe were probably the most growth-limiting nutrients. The growth rate of some diatoms was probably limited directly or indirectly at high pH. Laboratory biotests with natural populations may give valuable information on the growth-properties of different populations in the lakes. The biotests should, however, be carried out in combination with chemical and physical measurements and quantitative determinations of population densities.     

Defenses in phytoplankton against grazing induced by nutrient limitation, UV-B stress and infochemicals

It is becoming increasingly evident that the efficiency of zooplankton grazing on algae is not only a matter of quantity of the grazer relative to its food. Planktonic primary producers are not defenseless food-particles that are easily harvested by the consumers. Several algal species are able to adjust their phenotype (colony formation, spines, size) in such a way that it results in a reduced grazing pressure. It was recently demonstrated that morphological changes in the cell wall of green algae, induced by nutrient limitation and UV-B stress, may reduce their digestibility. A high fraction of induced cells pass intact and viable through the gut of the zooplankters, such that the grazing impact on the population is strongly reduced. It was also found that the presence of exudates (infochemicals) released by daphnids may change the morphology of algae. Unicellular green algae of the genus Scenedesmus were induced to form eight-cell coenobial types, heavily armed with spines, within three to five days after adding filtered water from an algal culture with Daphnia present. Both defence mechanisms may play an important role in zooplankton production and competition, and may serve as an example of highly efficient strategies to resist heavy grazing pressure.     

Inter-annual,seasonal and spatial variability in nutrient limitation of phytoplankton production in a river impoundment

We characterize seasonal and spatial patterns in phytoplankton abundance, production and nutrient limitation in a mesotrophic river impoundment located in the southeastern United States to assess variation arising from inter-annual differences in watershed inputs. Short-term (48 h) in situ nutrient addition experiments were conducted between May and October at three sites located along the longitudinal axis of the lake. Nutrient limitation was detected in 12 of the 18 experiments conducted over 2 years. Phytoplankton responded to additions of phosphorus alone although highest chlorophyll concentrations were observed in enclosures receiving combined (P and N) additions. Growth responses were greatest at downstream sites and in late summer suggesting that those populations experience more severe nutrient limitation. Interannual variation in nutrient limitation and primary production corresponded to differences in the timing of hydrologic inputs. Above average rainfall and discharge in late-summer (July–October) of 1996 coincided with higher in-lake nutrient concentrations, increased production, and minimal nutrient limitation. During the same period in 1995, discharge was lower, nutrient concentrations were lower, and nutrient limitation of phytoplankton production was more pronounced. Our results suggest that nutrient limitation is common in this river impoundment but that modest inter-annual variability in the timing of hydrologic inputs can substantially influence seasonal and spatial patterns.     

Temperature and nutrient availability interact to mediate growth and body stoichiometry in a detritivorous stream insect

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Nutrient limitation and stoichiometry of carnivorous plants

The cost-benefit model for the evolution of carnivorous plants posits a trade-off between photosynthetic costs associated with carnivorous structures and photosynthetic benefits accrued through additional nutrient acquisition. The model predicts that carnivory is expected to evolve if its marginal benefits exceed its marginal costs. Further, the model predicts that when nutrients are scarce but neither light nor water is limiting, carnivorous plants should have an energetic advantage in competition with non-carnivorous plants. Since the publication of the cost-benefit model over 20 years ago, marginal photosynthetic costs of carnivory have been demonstrated but marginal photosynthetic benefits have not. A review of published data and results of ongoing research show that nitrogen, phosphorus, and potassium often (co-)limit growth of carnivorous plants and that photosynthetic nutrient use efficiency is 20 - 50 % of that of non-carnivorous plants. Assessments of stoichiometric relationships among limiting nutrients, scaling of leaf mass with photosynthesis and nutrient content, and photosynthetic nutrient use efficiency all suggest that carnivorous plants are at an energetic disadvantage relative to non-carnivorous plants in similar habitats. Overall, current data support some of the predictions of the cost-benefit model, fail to support others, and still others remain untested and merit future research. Rather than being an optimal solution to an adaptive problem, botanical carnivory may represent a set of limited responses constrained by both phylogenetic history and environmental stress.     

Evidence that soil nutrient stoichiometry controls the competitive abilities of arbuscular mycorrhizal vs. root-borne non-mycorrhizal fungi

A majority of plant species has roots that are colonized by both arbuscular mycorrhizal (AM) and non-mycorrhizal (NM) fungi. The latter group may include plant mutualists, commensals, parasites and pathogens. The co-occurrence of these two broad groups may translate into competition for root volume as well as for plant-derived carbon (C). Here we provide evidence that the relative availability of soil nitrogen (N) and phosphorus (P) (i.e., soil nutrient stoichiometry) controls the competitive balance between these two fungal guilds. A decrease in the soil available N:P ratio resulted in a lower abundance of AM fungi and a corresponding increase in NM fungi. However, when the same fertilization treatments were applied in a soil in which AM fungi were absent, lowering the soil available N:P ratio did not affect NM fungal abundance. Taken collectively, our results suggest that the increase in NM fungal abundance was not a direct response to soil nutrient stoichiometry, but rather a competitive release from AM fungi responding negatively to higher soil P. We briefly discuss the mechanisms that may be responsible for this competitive release.