An in situ enclosure experiment to test the solar UVB impact on plankton in a high-altitude mountain lake. II. Effects on the microbial food web

We studied the impact of ambient levels of solar ultraviolet B (UVB) radiation on the planktonic microbial food web (viruses, heterotrophic bacteria, heterotrophic nanoflagellates and ciliates) of a high-mountain lake (2417 m above sea level) under in situ conditions for 16 days. Enclosures of 1 m³ receiving either the full sunlight spectrum or sunlight without UVB radiation were suspended at the lake surface. We found that the abundance of heterotrophic nanoflagellates was always lower in the +UVB treatment than in the -UVB one. In addition, bacterial consumption, measured by the disappearance of fluorescently labelled bacteria, was significantly (P < 0.05) reduced in the +UVB treatment. The abundance of non-filamentous bacteria (<10 m long) was also lower in the +UVB treatment, suggesting a direct effect of UVB on their growth. This was supported by the significantly (P < 0.05) lower cell-specific activity ([³H])thymidine incorporation) found on the fifth day of the experiment. In contrast, UVB radiation had no effect on filamentous bacteria (>1 m long), which represented only a small fraction of the total abundance (<4%), but up to 70% of the total bacterial biovolume. Ciliates, mainly Urotricha pelagica and Urotricha furcata, were less impacted by UVB radiation, although the net growth rate during the first week of the experiment was lower in the +UVB treatment than in the -UVB one (0.22 and 0.39 day^-1, respectively). The abundance of virus-like particles during the first week of the experiment was higher in the -UVB treatment. After reaching the maximum value for the interaction viruses x bacteria, viral abundance decreased dramatically (by 85%) in both treatments with a decay rate of 0.017 h^-1. This study illustrates the complexity in assessing the impact of UVB radiation when more than one trophic level is considered and indicates the existence of different sensitivity to UVB radiation among components of the microbial food web.      

围隔中鲢和菹草控藻效果及其生态学意义

在滇池沿岸的池塘中 ,构建了面积为 10 0m2 ,水深为 85cm ,底质均为淤泥的四个围隔 ,其中在三个围隔内分别放养密度为 10 0g/m3 、75g/m3 、5 0g/m3 的鲢。它们均在一定程度上控制浮游植物的生物量并改善水体透明度 ,其中初始放养鲢 75g/m3 的围隔控制效果最明显。放养三种密度的鲢在有效控制微囊藻水华的同时 ,原先水体的优势种类发生质的变化 ,水体中浮游植物的多样性也明显增加。放鱼 2个月后 ,在生物操纵的围隔中同时种入菹草 ,在75g/m3 的鲢控藻的围隔中 ,菹草长势良好 ,浮游动物大量出现 ,透明度进一步增加     

Analysis of phytoplankton-zooplankton relationships in an oligotrophic lake under natural and manipulated conditions

During August, 1987, we performed a series of Limnocorral experiments in lake La Caldera, a small winter-kill lake in which phytoplankton is strikingly nutrient-limited. The effects of biomanipulation on zooplankton-phytoplankton relationships were assessed by monitoring both individual species and whole-assemblage responses. Two sizes of enclosures were used (15 and 350 litres) and two treatments were assayed: 1) removal of zooplankton by 45 μm filter net and 2) doubling the natural grazing pressure by increasing the zooplankton concentration. Results show the two enclosure types to differ strikingly: flagellates disappeared from the small enclosures, resulting in four- to six-fold changes in chlorophylla concentration and three- to four-fold changes in number of individuals. Most species were grazed (a prey selectivity based on criteria other than size was observed) and their net growth rate increased with zooplankton concentration, causing a net increase in the phytoplankton growth, a stimulatory effect probably through nutrient regeneration that overrides the losses due to grazing.     

Effects of a filter-feeding fish [silver carp, Hypophthalmichthys molitrix (Val.)] on phyto- and zooplankton in a mesotrophic reservoir: results from an enclosure experiment

SUMMARY 1. Silver carp, Hypophthalmichthys molitrix (Val.), feeds on both phyto- and zooplankton and has been used in lake biomanipulation studies to suppress algal biomass. Because reports on the effects of silver carp on lake food webs have been contradictory, we conducted an enclosure experiment to test how a moderate biomass of the fish (10 g wet weight m⁻³) affects phytoplankton and crustacean zooplankton in a mesotrophic temperate reservoir.
2. Phytoplankton biomass <30 μm and particulate organic carbon (POC) <30 μm were significantly higher in enclosures with silver carp than in enclosures without fish, whereas Secchi depth was lower. Total copepod biomass declined strongly in both treatments during the experiment, but it was significantly higher in fish-free enclosures. Daphnid biomass was also consistently higher in enclosures without fish, although this effect was not significant. However, the presence of fish led to a fast and significant decrease in the size at maturity of Daphnia galeata Sars. Thus, the moderate biomass of silver carp had a stronger negative effect on cladoceran zooplankton than on phytoplankton.
3. Based on these results and those of previous studies, we conclude that silver carp should be used for biomanipulation only if the primary aim is to reduce nuisance blooms of large phytoplankton species (e.g. cyanobacteria) that cannot be effectively controlled by large herbivorous zooplankton. Therefore, stocking of silver carp appears to be most appropriate in tropical lakes that are highly productive and naturally lack large cladoceran zooplankton.     

Will reduced summer UV-B levels affect zooplankton populations of temperate humic and clearwater lakes?*

We collected zooplankton samples from 4×4 m enclosures located in two acidified lakes (pH<5) in Nova Scotia from June to August 1996. One lake had mean dissolved organic carbon concentrations greater than 10 mg l^–1 (brown water), while the other had values between 2 and 4 mg l^–1. In each lake, three enclosures were open to ambient light, while three were covered by Mylar® sheets which removed UVB and UVC wavelengths. Weekly sampling was done at all sites in July and near the end of August. Individuals were identified and the total community weighed. Analysis of results using both t-test and canonical analysis revealed small differences in populations between open and UVB covered clearwater sites in early July, but not later. There was no response of midsummer zooplankton communities in the humic lake which had a 95% extinction depth of 3 cm. In the clearwater lake (95% UVB extinction at 50 cm), the effect of radiation exclusion was relatively weak compared to most other published studies, but nevertheless statistically significant.     

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.     

Quantifying effects of phytoplankton on the heat budgets of two large limnetic enclosures

1. The aestival heat budgets of two large limnetic enclosures within a small lake in the English Lake District were studied. During summer, these enclosures had different nutrient supplies and consequently different phytoplankton populations. 2. As initial temperature profiles were similar and the incoming surface heat and momentum fluxes for the two enclosures were identical, subsequent changes in the heat budget were assumed to be induced by the biological differences between the enclosures. The proposed mechanism is an increased surface absorption of solar radiation leading to extra surface warming and a consequent excess loss of heat to the atmosphere through long‐wave emittance and sensible and latent heat fluxes, conservatively estimated to be of the order of 10–30 W m^?2. 3. Theoretical calculations show that potential effects on a heat budget could be considerably larger than those observed here. The inherent non‐linearity of the heat fluxes implies that such effects will be more important in warmer lakes than in colder ones. 4. Thermocline depth and strength were also altered by the response to differences in phytoplankton. 5. Any changes in climate or in nutrient loading from the catchment which substantially affect abundance or timing of phytoplankton populations in a lake will consequently also change the thermal structure of the lake.     

Negative effects of UVB-irradiated phytoplankton on life history traits and fitness of Daphnia magna

SUMMARY 1. We tested the effect of ultraviolet-B (UVB)-irradiated phytoplankton on life history characteristics of Daphnia magna . Two phytoplankton species were used, Chlamydomonas reinhardtii and Cryptomonas pyrenoidifera . The phytoplankton species were cultured under photosynthetically active radiation (PAR) conditions, and under PAR supplemented with ultraviolet-A and ultraviolet-B radiation, and fed to Daphnia .
2. Life history traits of Daphnia were negatively affected when fed on UVB-irradiated Cryptomonas . Size at maturity was depressed and fewer juveniles with lower fitness were produced in the UVB treatments. In the Chlamydomonas experiment, no significant effects were found.
3. The cause of the observed UVB effects is likely to be constraints in food quality. Ultraviolet-B radiation thus has the potential of inhibiting energy transfer from the first to the second trophic level.     

不同生活型大型植物对浮游植物群落的影响

湖泊加速富营养化是世界范围内的普遍现象,由此造成水质恶化,藻类大量增生,水生植被特别是沉水植物衰退乃至消失,生物多样性降低,严重影响湖泊主要功能的发挥。大型植物与浮游植物都是浅水湖泊的初级生产者,其间存在复杂的相互关系,如除竞争作用外,还可能存在相生...     

Ecosystem development in different types of littoral enclosures

Macrophyte growth was studied in two enclosure types (gauze and polythene) in a homogeneousPotamogeton pectinatus bed in Lake Veluwe (The Netherlands). The gauze was expected to allow for sufficient exchange with the lake to maintain similar seston densities, the polythene was expected to exclude fish activity and most water exchange. Polythene enclosures held higher totalP. pectinatus biomass (ash-free dry weight, AFDW) than the lake, gauze enclosures were intermediate. The enclosures had a higher abundance of other macrophyte species (Chara sp.,Potamogeton pusillus) than the lake. Seston ash content was not but seston AFDW, periphyton ash content and AFDW were lower in polythene than in gauze enclosures. The difference in plant biomass between gauze and polythene may be attributed to a difference in periphyton density and in seston AFDW due to zooplankton grazing (Rotatoria andDaphnia densities were higher in polythene enclosures). Since seston and periphyton AFDW and ash content were similar in lake and gauze enclosures, the intermediate macrophyte biomass in the gauze enclosures may be explained by reduced wave action and mechanical stress. Alternatively, phytoplankton inhibition by allelopathic excretions from the macrophytes may have caused the high macrophyte biomass in the polythene, and an absence of sediment-disturbing fish the intermediate biomass in the gauze enclosures. Creation of sheltered areas may favour macrophyte growth through both mechanisms and we conclude that this can be an important tool in littoral biomanipulation.     

Phytoplankton and zooplankton response to ultraviolet radiation in a high-altitude Andean lake: short- versus long-term effects

Exclusion experiments on global UV (A and B) radiation and globalUVB were performed in 460 I mesocosms with plankton communitiesfrom the oligotrophic Andean lake Laguna Negra (33°35'S–70°04'W;2700 m a.s.l.). The experiments were run for 30 days duringthe summers of 1991–1992 and 1992–1993, and for48 days in 1993–1994. When UVB radiation was allowed toenter into the mesocosms (full sun), the population of Ankyrajudayi (Chlorophyta) reached the highest density, suggestingthat this species can endure high levels of UV radiation. Concurrently,an increase in chlorophyll a concentration was observed in thistreatment. The cladoceran Chydorus sphaericus and the rotiferLepadella ovallts were strongly inhibited by UVB. Conversely,UVB radiation had no effect on the survival of the differentlife stages of the calanoid copepod Boeckela gractlipes, suggestinga species-specific difference in the sensitivity to solar UVBradiation. Moreover, no reduction in the number of copepod eggsper female and the number of nauplii produced was observed.Apparently, herbivory does not strongly affect phytoplanktonabundance. Moreover, the phytoplankton species composition changedin the different treatments over the time. Fragilaria construensand Fragilaria crotonensis were dominant in those mesocosmswhere UVB was excluded. Populations fluc tuated depending ontheir life cycles and the period of time they were exposed toUVB radiation. It is important to define the time scale of exclusionexperiments, because different conclusions about the influenceof UVB irradiance result from short-, medium- or long-term exposures.     

Bottom-up effects of bream (Abramis brama L.) in Lake Balaton

Enclosures (17 m³) were used in the mesotrophic area of Lake Balaton to determine the impact of benthivorous bream (Abramis brama L.) on the lower trophic levels during summers of 1984–86. In enclosures with a fish biomass similar to the biomass in the eutrophic area of the lake, the number of phytoplankton species was highest. In enclosures with a low fish biomass the phytoplankton was dominated by the greens. A high biomass of bream in the mesotrophic basin caused bacterial production corresponding to that of the eutrophic part of the lake. Crustaceans were dominated by copepods and were unable to control phytoplankton peaks. Bottom-up effects of bream were more obvious than top-down effects and seem to be more important in the possible control of water quality.     

Calanoid copepod grazing on phytoplankton: seasonal experiments on natural communities

Calanoid copepods are major components of most lacustrine ecosystems and their grazing activities may influence both phytoplankton biomass and species composition. To assess this we conducted four seasonal, in situ, grazing experiments in eutrophic Lake Rotomanuka, New Zealand. Ambient concentrations of late stage copepodites and adults of calanoid copepods (predominantly Calamoecia lucasi, but with small numbers of Boeckella delicata) were allowed to feed for nine days on natural phytoplankton assemblages suspended in the lake within 1160 litre polyethylene enclosures. The copepods reduced the total phytoplankton biomass of the dominant species in all experiments but were most effective in summer (the time of highest grazer biomass) followed by spring and autumn. In response to grazing pressure the density of individual algal species showed either no change or a decline. There were no taxa which increased in density in the presence of the copepods. The calanoid copepods suppressed the smallest phytoplankton species (especially those with GALD (Greatest Axial Linear Dimension) < µm) and there appeared to be no selection of algae on the basis of biovolume. Algal taxa which showed strong declines in abundance in the presence of the copepods include Cyclotella stelligera, Coelastrum spp., Trachelomonas spp., Cryptomonas spp., and Mallomonas akrokomos. Calanoid copepods are considered important grazers of phytoplankton biomass in this lake. The study supports the view that high phytoplankton:zooplankton biomass ratios and large average algal sizes characteristic of New Zealand lake plankton may, at least partly, be caused by year round grazing pressure on small algae shifting the competitive balance in favour of larger algal species.     

Modulation of the bacterial response to spectral solar radiation by algae and limiting nutrients

SUMMARY 1. The response of bacterial production (measured as [³H]TdR incorporation rate) to spectral solar radiation was quantified experimentally in an oligotrophic high-mountain lake over 2 years. Bacterial responses were consistent: ultraviolet-B (UVB) was harmful, whereas ultraviolet-A (UVA) + photosynthetically active radiation (PAR) and PAR enhanced bacterial activity. Full sunlight exerted a net stimulatory effect on bacterial activity in mid-summer but a net inhibitory effect towards the end of the ice-free period.
2. Experiments were undertaken to examine whether the bacterial response pattern depended on the presence of algae and/or was modulated by the availability of a limiting inorganic nutrient (phosphorus, P). In the absence of algae, [³H]TdR incorporation rates were significantly lower than when algae were present under all light treatments, and the consistent bacterial response was lost. This suggests that the bacterial response to spectral solar radiation depends on fresh-C released from algae, which determines the net stimulatory outcome of damage and repair in mid-summer.
3. In the absence of algae, UVB radiation inhibited bacteria when they were strongly P-deficient (mean values of N : P ratio: 46.1), whereas it exerted no direct effect on bacterial activity when they were not P-limited.
4. P-enrichment of lake water markedly altered the response of bacteria to spectral solar radiation at the end of ice-free period, when bacteria were strongly P-deficient. Phosphorus enrichment suppressed the inhibitory effect of full sunlight that was observed in October, both in whole lake water (i.e. including algae) and in the absence of algae. This indicates that the bacterial P-deficiency, measured as the cellular N : P ratio, was partly responsible for the net inhibitory effect of full sunlight, implying a high bacterial vulnerability to UVB.     

Studies on the Effects of Silver Carp (Hypophthalmichthys molitrix) on the Phytoplankton in a Shallow Hypereutrophic Lake through an Enclosure Experiment

The responses of nutrients, water transparency, zooplankton and phytoplankton to a gradient of silver carp biomass were assessed using enclosure methods. The gradient of four silver carp biomass levels was set as follows: 0, 116, 176 and 316 g m^—2. Nutrients did not show any statistically significant differences among the treatments. An outburst of Daphnia only occurred in fishless enclosures where phytoplankton biomass was the lowest and water clarity significantly increased. While among fish enclosures, the small‐sized Moina micrura dominated throughout the experiment and both zooplankton and phytoplankton biomasses decreased with increased fish biomass. No large colonial cyanobacterial blooms occurred in the fishless enclosures as predicted. This might be due to low water temperature, short experiment time and the occurrence of large bodied Daphnia in our experiment. Cryptophyta was the most dominant group in most of the enclosures and the lake water throughout the experiment. The fishless enclosure had much lower proportion of Cyanophyta but higher proportion of Trachelomonas sp.     

The effect of small zooplankton on the microbial loop and edible algae during a cyanobacterial bloom

SUMMARY 1. We studied the effect of the small crustacean zooplankton on heterotrophic micro-organisms and edible phytoplankton in a eutrophic lake during a cyanobacterial bloom.
2. Small (15 L) enclosures were filled with natural or screened (100 μm) lake water and incubated for 5 days in the lake. Screening removed crustacean zooplankton but the initial density of rotifers and phytoplankton remained the same in control and removal treatments. Changes in the abundance and biomass of bacteria, autotrophic picoplankton (APP), heterotrophic nanoflagellates (HNF) and ciliates were measured daily.
3. The crustacean zooplankton, dominated by the small cladoceran Chydorus sphaericus , did not affect cyanobacteria, the main phytoplankton group during the experiment.
4. The removal of the crustacean zooplankton induced a higher abundance of ciliates and reduced that of the HNF, indicating the importance of ciliates in controlling HNF in this system.     

Bioassay for phosphate demand in phytoplankton from acidified lakes: Lake Njupfatet,an example of phosphate deficiency induced by liming

A bioassay was developed, involving steady-state ATP level determinations, for estimation of phosphate demand and deficiency in natural phytoplankton communities. The studies were performed on phytoplankton from the moderately acidified Lake Njupfatet in central Sweden before and after liming. Phytoplankton samples from in situ enclosure experiments with low-dose enrichments of nitrate and phosphate and removal of large (> 100 µm) zooplankton and from the lake water were collected. The phytoplankton were concentrated by through-flow centrifugation and post-cultured in the laboratory with or without the addition of phosphate. A relative increase in the ATP:chlorophyll a ratio after the phosphate treatment as compared to samples without phosphate enrichment was found to be a highly reproducible indicator of phosphate deficiency in the natural phytoplankton population. In contrast, the absolute ATP:chlorophyll a ratio varied substantially between different sampling occasions. No phosphate deficiency was detected in phytoplankton from the acidic lake or from fertilized in situ enclosures. However, phytoplankton from in situ enclosures without added nutrients showed evidence of phosphate limitation after 21 days incubation. Also, the phytoplankton community developed a significant phosphate deficiency the summer after lake liming. The results from the ATP analyses are compared with chemical data of the lake water, phytoplankton community structure and phosphatase activities in the lake before and after liming. The average total biomass of phytoplankton and the average Tot-P measured during May to September decreased with appr. 30% after liming while Tot-N was essentially unaffected and the phosphatase activities increased by 1000–2000%.     

Effects of planktivorous and benthivorous fish on organisms and water chemistry in eutrophic lakes

The effects of planktivorous and benthivorous fish on benthic fauna, zooplankton, phytoplankton and water chemistry were studied experimentally in two eutrophic Swedish lakes using cylindrical enclosures. In enclosures in both lakes, dense fish populations resulted in low numbers of benthic fauna and planktonic cladocerans, high concentration of chlorophyll, blooms of blue-green, algae, high pH and low transparency. In the soft-water Lake Trummen, total phosporus increased in the enclosure with fish, but in the hard-water Lake Bysjön total phosphorus decreased simultaneously with precipitation of calcium carbonate. Enclosures without fish had a higher abundance of benthic fauna and large planktonic cladocerans, lower phytoplankton biomass, lower pH and higher transparency.The changes in enclosures with fish can be described as eutrophication, and those in enclosures without fish as oligotrophication. The possibility of regulation of fish populations as a lake restoration method is discussed.This paper was presented at the XXth SIL Congress in Copenhagen in 1977.     

Relative importance of periphyton and phytoplankton in turbid and clear vegetated shallow lakes from the Pampa Plain (Argentina): a comparative experimental study

We analyzed experimentally the relative contribution of phytoplankton and periphyton in two shallow lakes from the Pampa Plain (Argentina) that represent opposite scenarios according to the alternative states hypothesis for shallow lakes: a clear lake with submerged macrophytes, and a turbid lake with high phytoplankton biomass. To study the temporal changes of both microalgal communities under such contrasting conditions, we placed enclosures in the littoral zone of each lake, including natural phytoplankton and artificial substrata, half previously colonized by periphyton until a mature stage and half clean to analyze periphyton colonization. In the clear vegetated shallow lake, periphyton chlorophyll a concentrations were 3–6 times higher than those of the phytoplankton community. In contrast, phytoplankton chlorophyll a concentrations were 76–1,325 times higher than those of periphyton in the turbid lake. Here, under light limitation conditions, the colonization of the periphyton was significantly lower than in the clear lake. Our results indicate that in turbid shallow lakes, the light limitation caused by phytoplankton determines a low periphyton biomass dominated by heterotrophic components. In clear vegetated shallow lakes, where nitrogen limitation probably occurs, periphyton may develop higher biomass, most likely due to their higher efficiency in nutrient recycling.     

Effectiveness of phytoplankton control by large-bodied and small-bodied zooplankton

Employingin situ enclosures containing inocula of the lake zooplankton (mainlyDaphnia galeata, Daphnia cucullata andBosmina spp.) from a moderately eutrophic Lake Ros (Northern Poland) or large-bodiedDaphina magna, the following observations on succession of phytoplankton were made: 1) whereasD. magna could control the density of all the photoplankton size classes, the lake zooplankton could not suppress the large-sized phytoplankters or net phytoplankton; 2) the lake zooplankton was able to control the density of small algae (< 50μm), but its effect on large algae may be opposite: a promotion of net phytoplankton growth by removing small-sized algae which can out-compete net phytoplankton for limited PO₄-P resources (<5μg P l⁻¹). Since efficiency of phytoplankton density control byD. magna decreased with an increase in net phytoplankton abundance, biomanipulation could not be successful without introducing or maintaining a high population of large-bodied cladoceran species before high densities of large algae would make the control of phytoplankton inefficient.