Zooplankton response to shading effects of free-floating plants in shallow warm temperate lakes: a field mesocosm experiment

Dense mats of free floating plants (FFP) often produce severe underwater light attenuation and strong oxygen depletion in the water column. In this study, we experimentally assessed the zooplankton response to artificial shading using field mesocosms. During 30 days, we simulated three different light scenarios by mimicking the persistence, absence, and fluctuation of FFP typically encountered in vegetated shallow subtropical lakes. We used dark meshes to simulate the abiotic effects engineered by FFP. Both in the permanently covered and fluctuating situations, anoxia impaired zooplankton development. Anoxia constituted a major driving force in shaping the zooplankton response, whereas the feeding resource availability (phytoplankton) seemed to play a minor role; no top down effect on phytoplankton occurred in anoxic situations. In the fluctuating cover regime (periodic darkness and anoxia), the temporal variation of nanophytoplankton was not affected by zooplankton; once again oxygen availability seemed the main force shaping the zooplankton dynamics. Either periodical or permanent shading, associated to anoxic conditions, impaired the success of small herbivores. Large herbivores and microphytoplankton were negatively affected only under persistent shade and anoxia. In contrast, when neither light nor oxygen limitation occurred, such as in the scenario without shading, top-down control occurred. This study highlights the importance that the oxygen dynamics driven by the presence of FFP exert on the structure and dynamics of zooplankton assemblages and on the top down cascading effects on phytoplankton in warm temperate or subtropical shallow lakes.     

Size fractionated phytoplankton production in two humic shallow lakes with contrasting coverage of free floating plants

We conducted a 1-year survey in two humic shallow lakes from the floodplain of the Lower Paraná River, Laguna Grande Lake (LGL) and a relictual oxbow lake (ROL). We aimed to test two hypotheses: (1) the efficiency in light use of picoplankton (0.2–3 μm) is greater as light restriction increases and (2) the contribution of picoplankton to the total productivity is higher when the total photosynthetic biomass is lower. We performed P–E curves for picoplankton and nano- and microplankton (>3 μm) using the ¹⁴C assimilation technique. The light environments of the water bodies differed mainly owing to the development of free floating plants on the surface of the ROL and the dominance of phytoplankton in LGL. Primary productivity patterns in LGL were seasonality driven whilst in the ROL they were related to the coverage of floating macrophytes, which promoted light limitation and a lower productivity. In LGL, nano- and microplankton were in general more productive and the relative contribution of picoplankton to the total phytoplankton production decreased with the increase in total photosynthetic biomass. Hence, our study extends previously observed patterns to subtropical shallow lakes, where seasonality and free floating plants may influence the dynamics of phytoplankton production.     

Filamentous green algae inhibit phytoplankton with enhanced effects when lakes get warmer

1. Filamentous green algae (FGA) may represent an alternative state in high‐nutrient shallow temperate lakes. Furthermore, a clear water state is sometimes associated with the dominance of FGA; however, the mechanisms involved remain uncertain. 2. We hypothesised that FGA may promote a clear water state by directly suppressing phytoplankton growth, mostly via the release of allelochemicals, and that this interaction may be affected by temperature. 3. We examined the relationships between FGA, phytoplanktonic chlorophyll a concentrations and zooplankton in a series of mesocosms (2.8 m³) mimicking enriched shallow ponds now and in a future warmer climate (0 and c. 5 °C above ambient temperatures). We then tested the potential allelopathic effects of FGA (Cladophora sp. and Spirogyra sp.) on phytoplankton using several short‐term microcosms and laboratory experiments. 4. Mesocosms with FGA evidenced lower phytoplanktonic chlorophyll a concentrations than those without. Zooplankton and zooplankton : phytoplankton biomass ratios did not differ between mesocosms with and without FGA, suggesting that grazing was not responsible for the negative effects on phytoplanktonic biomass (chlorophyll a). 5. Our field microcosm experiments demonstrated that FGA strongly suppressed the growth of natural phytoplankton at non‐limiting nutrient conditions and regardless of phytoplankton initial concentrations or micronutrients addition. Furthermore, we found that the negative effect of FGA on phytoplankton growth increased up to 49% under high incubation temperatures. The experiment performed using FGA filtrates confirmed that the inhibitory effect of FGA on phytoplankton may be attributed to allelochemicals. 6. Our results suggest that FGA control of phytoplankton growth may be an important mechanism for stabilising clear water in shallow temperate lakes dominated by FGA and that FGA may play a larger role when lakes get warmer.     

The role of light for fish–zooplankton–phytoplankton interactions during winter in shallow lakes – a climate change perspective

1. Variations in the light regime can affect the availability and quality of food for zooplankton grazers as well as their exposure to fish predation. In northern lakes light is particularly low in winter and, with increasing warming, the northern limit of some present-day plankton communities may move further north and the plankton will thus receive less winter light.
2. We followed the changes in the biomass and community structure of zooplankton and phytoplankton in a clear and a turbid shallow lake during winter (November–March) in enclosures both with and without fish and with four different light treatments (100%, 55%, 7% and <1% of incoming light).
3. In both lakes total zooplankton biomass and chlorophyll- a were influenced by light availability and the presence of fish. Presence of fish irrespective of the light level led to low crustacean biomass, high rotifer biomass and changes in the life history of copepods. The strength of the fish effect on zooplankton biomass diminished with declining light and the effect of light was strongest in the presence of fish.
4. When fish were present, reduced light led to a shift from rotifers to calanoid copepods in the clear lake and from rotifers to cyclopoid copepods in the turbid lake. Light affected the phytoplankton biomass and, to a lesser extent, the phytoplankton community composition and size. However, the fish effect on phytoplankton was overall weak.
5. Our results from typical Danish shallow eutrophic lakes suggest that major changes in winter light conditions are needed in order to have a significant effect on the plankton community. The change in light occurring when such plankton communities move northwards in response to global warming will mostly be of modest importance for this lake type, at least for the rest of this century in an IPCC A2 scenario, while stronger effects may be observed in deep lakes.     

Top-down control of phytoplankton: the role of time scale, lake depth and trophic state

SUMMARY 1. One of the most controversial issues in biomanipulation research relates to the conditions required for top-down control to cascade down from piscivorous fish to phytoplankton. Numerous experiments have demonstrated that Phytoplankton biomass Top-Down Control (PTDC) occurs under the following conditions: (i) in short-term experiments, (ii) shallow lakes with macrophytes, and (iii) deep lakes of slightly eutrophic or mesotrophic state. Other experiments indicate that PTDC is unlikely in (iv) eutrophic or hypertrophic deep lakes unless severe light limitation occurs, and (v) all lakes characterised by extreme nutrient limitation (oligo to ultraoligotrophic lakes).
2. Key factors responsible for PTDC under conditions (i) to (iii) are time scales preventing the development of slow-growing inedible phytoplankton (i), shallow depth allowing macrophytes to become dominant primary producers (ii), and biomanipulation-induced reduction of phosphorus (P) availability for phytoplankton (iii).
3. Under conditions (iv) and (v), biomanipulation-induced reduction of P-availability might also occur but is insufficient to alter the epilimnetic P-content enough to initiate effective bottom-up control (P-limitation) of phytoplankton. In these cases, P-loading is much too high (iv) or P-content in the lake much too low (v) to initiate or enhance P-limitation of phytoplankton by a biomanipulation-induced reduction of P-availability. However, PTDC may exceptionally result under condition (iv) if high mixing depth and/or light attenuation cause severe light limitation of phytoplankton.
4. Recognition of the five different conditions reconciles previous seemingly contradictory results from biomanipulation experiments and provides a sound basis for successful application of biomanipulation as a tool for water management.     

Differential responses to UVR by bacterioplankton and phytoplankton from the surface and the base of the mixed layer

SUMMARY 1. We tested the influence of ultraviolet radiation (UVR) and shallow stratification on phytoplankton and bacterioplankton from the surface and the base of the mixed layer in two boreal lakes in north-western Ontario, Canada.
2. We measured phytoplankton biomass and production, bacterioplankton production and plankton respiration after transplantation under three solar radiation treatments: ambient radiation (Photosynthetically active radiation (PAR) + ultraviolet-A (UVA) + ultraviolet-B (UVB)), minus UVB (PAR + UVA) and PAR only. We repeated this experiment on three occasions in each lake during the summer.
3. Solar stress (measured as reduced growth and photoinhibition) was generally only found in the 'base phytoplankton' (i.e. originating from the base of the mixed layer). No inhibition of photosynthesis by UVB exposure was found in near-surface phytoplankton. On the other hand, production of near-surface bacterioplankton was reduced following a 4-h UVR exposure but had increased after a 48-h exposure to both UVA and UVB compared with the PAR only treatment.
4. Negative effects of UVR on phytoplankton and bacterioplankton were not ubiquitous. We emphasise the importance of conducting experiments repeatedly, particularly those which test the effects of UVR on different community assemblages from different lakes.     

The impact of free-floating plant cover on phytoplankton assemblages of oxbow lakes (The Bug River Valley, Poland)

In the present study we focused on the impact of macrophyte cover (composed mainly of the Lemna genus) on phytoplankton taxonomic and functional diversity. Some important environmental parameters, mainly light (Kd_PAR), and the chemical conditions (pH, dissolved oxygen, ammonium, soluble and total forms of phosphorus) were closely related to the pleustophyte cover. Among them, the key factor in the phytoplankton ecology of the studied oxbow lakes was the dense macrophyte cover which strongly reduced the illumination of water. Neither differences in the mean nutrient concentrations between the lakes with FFP (Free Floating Plants) absence and those with FFP dominance nor significant relationships between nutrients and the phytoplankton structure were observed. The species composition of phytoplankton and the functional (FG) and morpho-functional (MFG) groups reflected the differences between the habitats connected with hydromacrophytes. The free-floating macrophyte cover favours mixotrophic and heterotrophic species, mainly Euglenophyta (coda W1 and W2) and chrysophytes (codon Ws) as well as shade-adapted cyanobacteria with the high tolerance of the low oxygen content (codon K). In lakes with FFP absence — taxa from Chlorophyta and Bacillariophyceae (associations X1, J, and D), or filamentous cyanobacteria (codon S1) dominated the phytoplankton. MFG were less related to the oxbow type and exhibited greater similarity between lakes independently of the presence or absence of FFP. Only unicellular Cyanoprokaryota which created MFG 4 and colonial chroococcales (MFG 5b and 5c) reached a greater percentage share, especially in oxbow lakes with FFP dominance.     

Responses of phytoplankton and epilithon during acidification and early recovery of a lake

1. Lake 302S in the Experimental Lakes Area of Canada was acidified from pH 6.7 (1981) to 5.1 (1986). The pH was further reduced to 4.5 in 1987 and held at that level until 1991. From 1992 to 1995, the pH was allowed to increase to a target value of 5.8.
2. The response of the phytoplankton community to decreasing pH from 6.0 to 5.1 was similar to that observed in another experimentally acidified lake (223) and in other atmospherically acidified lakes. Acidification affected species diversity of both the phytoplankton and epilithon. Phytoplankton diversity was positively correlated with pH. Epilithic algal diversity was more variable and did not correlate with pH.
3. Phytoplankton biomass was enhanced by acidification as the assemblage shifted from a dominance of chrysophytes to large dinoflagellates ( Gymnodinium sp. and Peridinium inconspicuum ). Epilithon biomass was unaffected, but dominance shifted from filamentous cyanophytes ( Lyngbya ) to acidophilic diatoms ( Tabellaria quadriseptata and Anomoeonis brachysira ).
4. The only taxon to be similarly affected in both the phytoplankton and epilithon was the cyanobacteria, being significantly reduced below pH 5.1. During early recovery (pH 5.5–5.8), cyanobacteria increased and species present prior to acidification recolonized both habitats.
5. In the early stages of recovery, planktonic and benthic assemblages remained more similar to acidified than natural assemblages, but more profound change began at pH > 5.5.     

在太湖中栽种沉水植物能使水变清吗？

浅水湖泊富营养化修复是环境科学领域的前沿研究.在富营养化的太湖中栽种沉水植物能使水变清,进而"向后跃变"为"草型湖泊"吗?回答是几乎不可能.原因在于藻类和沉水植物之间存在着竞争.在湖泊中决定沉水植物能否生长的关键是水体的光照条件.在富营养化的湖泊中,藻类对沉水植物有明显的"遮光效应",从而降低了它的竞争能力.另一种情况,枝角类的消失或者数量减少对藻类有利."下行效应"在浅水湖泊中可能更为重要,因此浅水湖泊中滤食性鱼类对浮游动物的捕食控制可能更强,导致枝角类几乎不可能通过牧食控制藻类的生长.在这样的状况下,沉水植物的恢复几乎是不可能的.如果沉水植物再遭受其它的损害或者恶劣的天气,就会突然一下子全部消失.因此在富营养化的湖泊中栽种沉水植物的成活率是很低的.这与浅水湖泊二种替代性稳定状态概念是相一致的.     

The effects of disturbance events on phytoplankton community structure in a small temperate reservoir

1 The effects of disturbances, in the form of storm events, on phytoplankton community structure were examined over the course of four years in Eau Galle Reservoir, Wisconsin, USA.
2 Disturbances consistently brought about significant, but highly transient, increases in apparent phytoplankton species richness. It is likely that these resulted from temporary increases in the biomass of previously undetected rare species.
3 Substantial shifts in community dominance were confined to large, early season events, and were seldom long-lived. Later 'climax' communities were highly resistant to any changes in dominance, even when increases in species richness occurred.
4 Regardless of when they occurred, disturbances tended to favour species from a narrow range of the successional sequence.     

Macrophytes moderate the taxonomic and functional composition of phytoplankton assemblages during a nutrient loading experiment

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Seasonal dynamics of macrophytes and phytoplankton in shallow lakes: a eutrophication‐driven pathway from plants to plankton?

1. Seasonal relationships between macrophyte and phytoplankton populations may alter considerably as lakes undergo eutrophication. Understanding of these changes may be key to the interpretation of ecological processes operating over longer (decadal‐centennial) timescales. 2. We explore the seasonal dynamics of macrophytes (measured twice in June and August) and phytoplankton (measured monthly May–September) populations in 39 shallow lakes (29 in the U.K. and 10 in Denmark) covering broad gradients for nutrients and plant abundance. 3. Three site groups were identified based on macrophyte seasonality; 16 lakes where macrophyte abundance was perennially low and the water generally turbid (‘turbid lakes’); 7 where macrophyte abundance was high in June but low in August (‘crashing’ lakes); and 12 where macrophyte abundance was high in both June and August (‘stable’ lakes). The seasonal behaviour of the crashing and turbid lakes was extremely similar with a consistent increase in nutrient concentrations and chlorophyll‐a over May–September. By contrast in the stable lakes, seasonal changes were dampened with chlorophyll‐a consistently low (<10–15 μg L^?1) over the entire summer. The crashing lakes were dominated by one or a combination of Potamogeton pusillus, Potamogeton pectinatus and Zannichellia palustris, whereas Ceratophyllum demersum and Chara spp. were more abundant in the stable lakes. 4. A long‐term loss of macrophyte species diversity has occurred in many shallow lakes affected by eutrophication. One common pathway is from a species‐rich plant community with charophytes to a species‐poor community dominated by P. pusillus, P. pectinatus and Z. palustris. Such compositional changes may often be accompanied by a substantial reduction in the seasonal duration of plant dominance and a greater tendency for incursions by phytoplankton. We hypothesise a slow‐enacting (10–100 s years) feedback loop in nutrient‐enriched shallow lakes whereby increases in algal abundance are associated with losses of macrophyte species and hence different plant seasonal strategies. In turn such changes may favour increased phytoplankton production thus placing further pressure on remaining macrophytes. This study blurs the distinction between so‐called turbid phytoplankton‐dominated and clear plant‐dominated shallow lakes and suggests that plant loss from them may be a gradual process.     

Ecosystem-level effects of re-oligotrophication and N:P imbalances in rivers and estuaries on a global scale

Trends and ecological consequences of phosphorus (P) decline and increasing nitrogen (N) to phosphorus (N:P) ratios in rivers and estuaries are reviewed and discussed. Results suggest that re-oligotrophication is a dominant trend in rivers and estuaries of high-income countries in the last two–three decades, while in low-income countries widespread eutrophication occurs. The decline in P is well documented in hundreds of rivers of United States and the European Union, but the biotic response of rivers and estuaries besides phytoplankton decline such as trends in phytoplankton composition, changes in primary production, ecosystem shifts, cascading effects, changes in ecosystem metabolism, etc., have not been sufficiently monitored and investigated, neither the effects of N:P imbalance. N:P imbalance has significant ecological effects that need to be further investigated. There is a growing number of cases in which phytoplankton biomass have been shown to decrease due to re-oligotrophication, but the potential regime shift from phytoplankton to macrophyte dominance described in shallow lakes has been documented only in a few rivers and estuaries yet. The main reasons why regime shifts are rarely described in rivers and estuaries are, from one hand the scarcity of data on macrophyte cover trends, and from the other hand physical factors such as peak flows or high turbidity that could prevent a general spread of submerged macrophytes as observed in shallow lakes. Moreover, re-oligotrophication effects on rivers may be different compared to lakes (e.g., lower dominance of macrophytes) or estuaries (e.g., limitation of primary production by N instead of P) or may be dependent on river/estuary type. We conclude that river and estuary re-oligotrophication effects are complex, diverse and still little known, and in some cases are equivalent to those described in shallow lakes, but the regime shift is more likely to occur in mid to high-order rivers and shallow estuaries.     

Effects of plant harvesting and nutrient enrichment on phytoplankton community structure in a shallow urban lake

The utility of shallow water bodies in urban environments is frequently compromised either by dense beds of submerged plants or cyanobacterial blooms associated with nutrient enrichment. Although submerged plants are often harvested to facilitate recreational uses, this activity may alter the phytoplankton community, which in turn, also may restrict the use of the lake. We tested whether (i) plant harvesting reduced the abundance of flagellate algae and increased the abundance of cyanobacteria, and (ii) whether increasing levels of nutrient enrichment caused shifts in the dominance of heterocytous cyanobacteria, non-heterocytous cyanobacteria and Chlorophyta, in a shallow urban lake in Southern Australia as has been observed for shallow Danish lakes in previous studies. These predictions were tested with large (3000 l), replicated mesocosms in a warm, highly productive, shallow lake densely colonised by the submerged angiosperm, Vallisnaria americana Michaux. The heterokont algae, Chlorophyta, Cyanobacteria and Cryptophyta were the most numerous algal divisions in the lake. The Euglenophyta, although uncommon in early summer, became more abundant towards the end of summer. The Dinophyta and Charophyta were rare. The abundance of the heterokont algae and Euglenophyta was significantly reduced by plant harvesting even after plants had partially re-established 18 weeks after initial harvesting. The decline in the Euglenophyta in response to plant harvesting is consistent with earlier findings, that the relative abundance of flagellate algae tends to be greater in the presence of submerged plants. Contrary to our prediction, we found that the Cyanobacteria did not increase in response to plant harvesting, however the response may be altered under higher nutrient levels. Algal responses to nutrient enrichment in the presence of dense V. americana plants generally followed the patterns observed in shallow Danish lakes despite the large differences in climatic conditions. Both studies found that the abundance of heterocytous cyanobacteria declined at higher levels of nutrient enrichment, whereas non-heterocytous cyanobacteria and chlorophytes increased.     

模拟水流条件下初级生产力及光动力学参数

为探讨水动力作用及其他物理因子改变对湖泊初级生产力的影响 ,1999年 5月 8日～ 6月 2 4日在中国科学院太湖湖泊生态系统研究站大型生态实验槽内进行模拟水动力实验 ,分 3种水流状态 2种光强测定初级生产力及其他相关参数。分析了初级生产力、光合速率的垂直分布 ,光合速率随光强的变化 ,并借助光动力学模型拟合得到光动力学参数。结果表明 ,在静止状态下 ,当水表面光强大于 5 0 0μmol/ (m2 · s)时 ,0～ 0 .4 m处存在光抑制现象 ,最大初级生产力出现在 0 .4～ 0 .6 m,此后由于动力作用使水体悬浮物增加 ,改变了水下光照条件 ,致使最大初级生产力呈向上移动的趋势 ,出现在 0～ 0 .2 m间 ;光合速率在静止状态下随深度递减缓慢 ,而到大水流状态则递减极为迅速 ,大水流状态下的平均光合速率明显低于静止状态和小水流状态 ;基于 2种类型的光动力学模型进行非线性拟合得到的 P- I曲线相关性很好 ,2种模型模拟的结果比较接近 ,基本上能够反映太湖光合速率随光强变化的实际情况 ;在太湖这种大型浅水湖泊 ,水动力的作用使得水体中悬浮物增加 ,造成光强的迅速衰减 ,这可能会大大降低湖泊的初级生产过程     

Lime treatment and its effects on the chemistry and biota of hardwater eutrophic lakes

1. The main focus of this study was to investigate the effects of single and multiple moderate doses of lime (slaked lime, Ca(OH)₂, and/or calcite, CaCO₃) on eutrophic hardwater lakes. This information would contribute to strategies to manage phytoplankton and macrophyte biomass in eutrophic lakes.
2. Water chemistry and biota were monitored for up to 7 years after initial lime treatment and results were compared with reference systems.
3. Complementary studies investigated the effect of lime on macrophytes in ponds, irrigation canals and microcosm experiments.
4. When water pH was kept within its natural range (≤ 10), single and multiple lime applications to lakes and ponds controlled macrophyte biomass, without negatively affecting invertebrate communities.
5. Single lime treatments at moderate dosages of lakes and ponds resulted in variable and mostly temporary changes in chlorophyll a (chl a ) and phosphorus (P) concentration. Although sediment P release was reduced in single-dose lakes during the first winter following treatment, reductions appeared temporary.
6. Multiple treatments of lakes and ponds were effective at reducing both chl a and P concentrations over longer periods. Mean winter P release rate was also reduced after initial treatment.
7. In laboratory studies, sediment cores were incubated with eight different treatments to assess P release. Redox-sensitive treatments were no more effective at lowering total P concentration in overlying water than some redox-insensitive treatments. Lime reduced total P concentrations, but was not as effective as treatments with alum.
8. The use of lime in managing macrophyte and phytoplankton biomass in shallow, hardwater lakes and ponds may be preferable over other treatments, because lime is economical and non-toxic as long as pH is kept within a natural range.     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Size-fractionated phytoplankton biomass and its implications for the dynamics of an oligotrophic tropical lake

1. Size-fractionated phytoplankton biomass was examined in relation to the hydrodynamics of tropical Lake Alchichica from 1999 to 2002.
2. Alchichica is a warm monomictic lake, in which mixing takes place from late December to early March. The lake is oligotrophic (mean total chlorophyll- a concentration 4.2 ± 4.2  μ g L⁻¹) and its phytoplankton biomass is dominated (72.3 ± 16.4%) by large individuals (>2  μ m). The degree of dominance of the large size class (nano- and microplankton) over the small size class (picoplankton) throughout the year is mainly determined by the availability of silicate and the Si/N ratio in the hypolimnion prior to the mixing period.
3. This is the first record of an oligotrophic tropical lake dominated by large size fractions of phytoplankton. Because of this dominance, the fate of most primary productivity is rapid sedimentation to the bottom followed by decomposition that promotes an anoxic hypolimnion.
4. Our findings in tropical Lake Alchichica challenge the idea that oligotrophic waters are dominated by small phytoplankton, as has been well established for the oligotrophic ocean and temperate lakes.     

Experimental studies of zooplankton–phytoplankton–nutrient interactions in a large subtropical lake (Lake Okeechobee, Florida, U.S.A.)

1. Over a 1-year period, twenty controlled experiments were performed using small mesocosms (20-l clear plastic carboys) and plankton communities collected from four sites in shallow, subtropical Lake Okeechobee, Florida. In replicated treatments, macrozooplankton grazers were excluded by size fractionation (115 μm), and/or nutrients (N and P) were added, and impacts on phytoplankton biomass and productivity were measured after 3-day incubations.
2. In most experiments (fifteen out of twenty), there was no significant effect of zooplankton exclusion on phytoplankton biomass or productivity, but there were significant increases in those attributes due to nutrient additions. The magnitude of the responses was a function of light availability at the collection sites.
3. In three experiments, zooplankton exclusion led to declines in phytoplankton biomass and productivity, suggesting that animals may sometimes have net positive effects on the phytoplankton, perhaps via nutrient recycling.
4. In only two experiments was there evidence of net negative impacts of grazers on the phytoplankton. In both instances, cladocerans ( Daphnia ambigua and Eubosmina tubicen ) were dominant in the zooplankton. However, the increases in chlorophyll a due to zooplankton exclusion were small (5–20%), probably because of the small size and relatively low grazing rates of the cladocerans.
5. The results support the hypothesis that phytoplankton biomass in Lake Okeechobee is little affected by herbivorous macrozooplankton. This may be a common feature of lowland tropical and subtropical lakes.     

Relationships between picophytoplankton and environmental variables in lakes along a gradient of water colour and nutrient content

SUMMARY 1. Biomass and production of picophytoplankton, phytoplankton and heterotrophic bacterioplankton were measured in seven lakes, exhibiting a broad range in water colour because of humic substances. The aim of the study was to identify environmental variables explaining the absolute and relative importance of picophytoplankton. In addition, two dystrophic lakes were fertilised with inorganic phosphorus and nitrogen, to test eventual nutrient limitation of picophytoplankton in these systems.
2. Picophytoplankton biomass and production were highest in lakes with low concentrations of dissolved organic carbon (DOC), and DOC proved the factor explaining most variation in picophytoplankton biomass and production. The relationship between picophytoplankton and lake trophy was negative, most likely because much P was bound in humic complexes. Picophytoplankton biomass decreased after the additions of P and N.
3. Compared with heterotrophic bacterioplankton, picophytoplankton were most successful at the clearwater end of the lake water colour gradient. Phytoplankton dominated over heterotrophic bacteria in the clearwater systems possibly because heterotrophic bacteria in such lakes are dependent on organic carbon produced by phytoplankton.
4. Compared with other phytoplankton, picophytoplankton did best at intermediate DOC concentrations; flagellates dominated in the humic lakes and large autotrophic phytoplankton in the clearwater lakes.
5. Picophytoplankton were not better competitors than large phytoplankton in situations when heterotrophic bacteria had access to a non-algal carbon source. Neither did their small size lead to picophytoplankton dominance over large phytoplankton in the clearwater lakes. Possible reasons include the ability of larger phytoplankton to float or swim to reduce sedimentation losses and to acquire nutrients by phagotrophy.