A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

The importance of Protozoa in controlling the abundance of planktonic algae in lakes

Apart from ciliates, there are other protozoans which can form an important element in the herbivorous zooplankton of lakes.
The protozoans discussed in this paper feed on colonial algae. They can greatly reduce the numbers of certain Chlorophyta in Windermere, Esthwaite Water and Blelham Tarn, lakes in the English Lake District.
A very short period, often 7–14 days, can suffice for the destruction of more than 99% of an algal population.
Experiments with a species of Pseudospora showed that it would only ingest one out of six green algal species, although, in nature, Pseudosporae which are morphologically similar to this species have been observed in all these algae.
The importance of such protozoans, and of parasitic fungi, has been underrated or neglected in studies of the quantitative relationships between primary and secondary production in the plankton.     

The relative importance of sediment and water column supplies of nutrients to the growth and tissue nutrient content of the green macroalga Enteromorpha intestinalis along an estuarine resource gradient

Large blooms of opportunistic green macroalgae such as Enteromorpha intestinalis are of ecological concern in estuaries worldwide. Macroalgae derive their nutrients from the water column but estuarine sediments may also be an important nutrient source. We hypothesized that the importance of these nutrient sources to E. intestinalis varies along a nutrient-resource gradient within an estuary. We tested this in experimental units constructed with water and sediments collected from 3 sites in Upper Newport Bay estuary, California, US, that varied greatly in water column nutrient concentrations. For each site there were three treatments: sediments + water; sediments + water + Enteromorpha intestinalis (algae); inert sand + water + algae. Water in units was exchanged weekly simulating low turnover characteristic of poorly flushed estuaries. The importance of the water column versus sediments as a source of nutrients to E. intestinalis varied with the magnitude of the different sources. When initial water column levels of dissolved inorganic nitrogen (DIN) and soluble reactive phosphorus (SRP) were low, estuarine sediments increased E. intestinalis growth and tissue nutrient content. In experimental units from sites where initial water column DIN was high, there was no effect of estuarine sediments on E. intestinalis growth or tissue N content. Salinity, however, was low in these units and may have inhibited growth. E. intestinalis growth and tissue P content were highest in units from the site with highest initial sediment nutrient content. Water column DIN was depleted each week of the experiment. Thus, the water column was a primary source of nutrients to the algae when water column nutrient supply was high, and the sediments supplemented nutrient supply to the algae when water column nutrient sources were low. Depletion of water column DIN in sediment + water units indicated that the sediments may have acted as a nutrient sink in the absence of macroalgae. Our data provide direct experimental evidence that macroalgae utilize and ecologically benefit from nutrients stored in estuarine sediments.     

Effects of Phosphorus Nutrient on the Hydraulic Conductivity of Sorghum （Sorghum vulgare Pers.） Seedling Roots Under Water Deficiency

Hydroponic experiments were conducted in a growth chamber and changes in the hydraulic conductivity of sorghum (Sorghum vulgare Pers.) roots (Lpr) at the three-leaf stage were measured using the pressure chamber method. Water deficiency was imposed with polyethylene glycol (PEG) 6000 and the phosphorus (P) levels were controlled by complete Hoagland solution with and without P nutrient. The objective of this study was to investigate the effect of P nutrition on root Lpr under water deficiency. The results showed that the Lpr in P deficiency treatments decreased markedly, but the Lpr recovered to the same value as that of control when sufficient P was supplied for 4-24 h. Water deficiency decreased Lpr, but the hydraulic conductivity of the roots with sufficient P supply was still higher than that of plants without P supply. When resuming water supply, the Lpr of the water-deficient plants under P supply recovered faster than that of plants without P supply, which indicates that plants with sufficient P nutrient are more drought tolerant and have a greater ability to recover after drought. The treatment of HgCl2 indicated that P nutrient could regulate the Lpr by affecting the activity and the expression levels of aquaporins.     

Factors that Control the Species Composition of Freshwater Phytoplankton,with Special Attention to Nutrient Concentrations

Variations of species composition and population size of planktonic algae were studied in relation to the different nutrient levels in a eutrophic pond and an oligotrophic lake. The results obtained were discussed in comparison with the changes in photosynthesis and chlorophyll of several algal populations cultured under different nutrient conditions. As based on unialgal culture experiments two types of algae (the Chlorella-type and the Melosira-type) could be distinguished with regard to variations of photosynthesis and chlorophyll in response to different nutrient levels. Distribution of the Chlorella-type algae may be confined to eutrophic waters, while the Melosira-type algae can be distributed widely in waters from oligotrophic to eutrophic.     

Scraping a living: a review of littorinid grazing

Littorinid snails are predominantly herbivorous and the versatility of their radulae enables them to feed on a variety of macroscopic and microscopic plants in a diversity of habitats. Some are selective feeders preferring some species of algae to others, and rejecting some even after a prolonged period of starvation. Different species of snail exhibit different preferences. The factors affecting the attractiveness and edibility of food plants are discussed and food value considered.Foraging behaviour of littorinids is briefly reviewed in relation to the influence of chemical cues from the algae. Littorinids appear to be able to select or reject algae without having ingested them, having perceived the plants from a distance, moving towards favoured foods (or habitat-providing plants) and away from those that it rejects. The nature of the chemical cues emitted by the algae is discussed. Temporal patterns of foraging activity show some evidence of an endogenous component which can be overridden by responses to environmental conditions. These patterns place restraints on energy intake.The structural and chemical defences used by algae against littorinid grazing are considered. The importance of polyphenolic compounds is evaluated. The effects of grazing as a selective agency and a factor influencing algal populations are discussed. There is some evidence that life history patterns are a response to grazing. The influence of external physical factors, such as salinity on grazing pressure is demonstrated.Finally, the impact of littorinid snails on intertidal communities is assessed in relation to their abundance and biogeographical distribution. The relative importance of littorinids is contrasted on shores possessing or lacking limpets.     

Flood disturbance, algal productivity, and interannual variation in food chain length

The length of a river food chain changed from year to year, shifting with the hydrologic regime. During drought years, grazers suppressed algae across a nutrient gradient, while predators were functionally unimportant. Following flood disturbance, predators suppressed grazers, releasing algae. These results suggest that hydrologic regime, rather than productivity, determines the functional length of this river food chain. Within years, algae and grazer biomass responded to an experimental productivity gradient in patterns predicted by simple trophic models that assume efficient energy transfer. Understanding differences among species within trophic levels, however, was crucial in delineating the controlling interactions.     

Filamentous green algae in freshwater streams on Signy Island,Antarctica

The streams of Signy Island are varied and extremely seasonal environments. Water flows from November/December to March/April; streams are frozen for the rest of the year. Streams usually flow through small, barren catchments and are nutrient poor, though they may be enriched by dense summer populations of seabirds and seals. Temperatures are consistently low. Stream depth is maximal during the spring melt period, declining over the course of the summer. Vegetation is exclusively algal, and filamentous chlorophytes from a particularly conspicuous component. Small numbers of vegetative cells survive the long frozen period in situ. A steady increase in standing crop results in a maximum 2 to 3 months after flow begins. Sloughing is the major loss mechanism and grazers are effectively absent. Three taxa of filamentous algae are common in Signy streams, species of Zygnema, Mougeotia and Klebsormidium. The distributions of these algae are described and related to physical and chemical features of their environment.     

The effect of benthic algae on phosphorus exchange between sediment and overlying water in shallow lakes: a microcosm study using 32P as a tracer

Sediments are of key importance in determining the nutrient levels of water in shallow lakes as they can act as either source or sink for phosphorus (P) depending on environmental conditions, sediment characteristics, and external nutrient loading. We examined the role of benthic algae in the P cycling between sediment and overlying water in experiments using ³²P as a tracer. Sediment and water samples were collected from Huizhou West Lake, a shallow, eutrophic waterbody located in Huizhou City, South China. Laboratory cultured benthic algae were transferred to cover the sediment core in tubes. When ³²P was added to the water in experimental tubes containing sediment cores with and without benthic algae, ³²P activity after 48 h was significantly lower in the tubes with algae, indicating that benthic algae removed P from the overlying water. When the tracer was injected into the sediment, ³²P activity in the water overlying sediment with benthic algae was substantially lower than in tubes with naked sediment, suggesting that benthic algae reduce the release of sediment P. Oxygen levels were significantly higher in the upper 3 mm of the sediments covered by benthic algae; thus, we hypothesized that oxygen produced by the algae helps inhibit the release of P from the sediment. Our study demonstrates that benthic algae are capable of reducing P levels in water overlying the sediment, suggesting that loss of benthic algae during eutrophication triggered by impoverished light conditions may accelerate the shift in shallow lakes from a clear water to a turbid state.     

Drought survival,summer dormancy and dehydrin accumulation in contrasting cultivars of Dactylis glomerata

To study survival under prolonged and severe drought in the perennial grass Dactylis glomerata we compared dormant, resistant and sensitive cultivars (cvs.) in both field and glasshouse experiments. Water status, membrane stability and expression of dehydrins were assessed in the immature leaf bases, which are the last surviving organs. Analysis of leaf elongation and senescence of aerial tissues showed that dormancy was exhibited by the potentially dormant cultivar (cv.) only in the field. This cultivar exhibited a high survival rate, similar levels of dehydration and expression of a low-molecular weight (22–24 kDa) dehydrin in both drought and irrigated plants, whether fully dormant or not. At the same level of soil water deficit, there were no differences between the non-dormant drought resistant and drought sensitive cultivars in plant water status and membrane stability. However, the accumulation of dehydrins as drought progressed was markedly different between these cultivars and was associated with their contrasting survival. The possible role of the major low-molecular dehydrins in maintenance of cell integrity under dehydration is discussed with reference to both summer dormancy and survival under severe drought.     

Eutrophication-induced changes in benthic algae affect the behaviour and fitness of the marine amphipod Gammarus locusta

This study, conducted in mesocosms, natural field sites, and in laboratory aquaria, showed that eutrophication altered the nutrient status and dominance patterns among marine macroalgae, which in turn, stimulated gammaridean density. Gammaridean abundance correlated positively with both nutrient addition and the amount of green algae (also stimulated by nutrient enrichment). Path analysis indicated that the direct effect of nutrients on gammaridean density was of less importance than the indirect effect through increased production of green algae. In cage colonisation experiments, either in the field or in a control mesocosm kept under ambient nutrient conditions, more gammarids colonised nutrient enriched algae (E-algae) than algae with ambient nutrient levels (A-algae). Gammarus locusta generally grew faster on nutrient enriched algal specimens and when reared on green rather than on brown algae (fucoids). The nutrient status of periphytic algae did not affect gammaridean growth significantly, but the number of egg-carrying females (and thus egg production) was significantly higher among gammarids reared on E-periphyton. The gammaridean habitat preference order (red > green > brown > periphyton) was almost the reverse of their growth rate in feeding assays (periphyton > green > brown). This implies that macroalgae may be more important as a habitat than as a food source for these animals, which then have to become mobile in search of optimal food items. In this process, algal nutrient content was important as the gammarids in our study actively chose high quality nutrient-rich food, which, in addition, increased their fitness. Stimulated growth rates and egg production may ultimately lead to population increase, which, combined with the preference for high nutrient food items may dampen the initial effect of nutrient enrichment (i.e. blooms of green macroalgae) in shallow coastal waters.     

The effect of a lake fertilization on the stability and material utilization of a limnetic ecosystem

The interaction between the phytoplankton, zooplankton and fish populations and certain abiotic environmental factors, was investigated in an oligotrophic Norwegian lake during a 5-yr period (1974–1978). The effects of adding artificial fertilizer in 1975 and 1976 were also studied. When cladoceran dominated, the zooplankton community was able to maintain a more or less constant phytoplankton biomass and a rather low phytoplankton production even when nutrient levels were raised. In years when rotifers were dominant, algal biomass and productivity increased, despite the amount of added nutrients being lower. The regression for the relationship between daily phytoplankton P/B and daily herbivore zooplankton P/B indicated that these trophic levels were highly interdependent. A change, from large-sized to smaller herbivorous zooplankton, due to fish predation, also led to an increase in phytoplankton turnover. The investigations show that planktivorous fish may be the key factor which determines the stability of limnetic systems and controls the material transfer from the algae to the higher trophic level.     

Experimental manipulation of water levels in two French riverine grassland soils

In this experimental study, we simulated the effects of different river flooding regimes on soil nutrient availability, decomposition and plant production in floodplain grasslands. This was done to investigate the influences of soil water contents on nutrient cycling. Water levels were manipulated in mesocosms with intact soil turfs from two French floodplain grasslands. Three water levels were established: a `wet' (water level at the soil surface), an `intermediate' (water level at –20 cm) and a `dry' treatment (water level at –120 cm). With increasing soil moisture, soil pH became more neutral, while redox-potential and oxygen concentration decreased. The `dry' treatment showed much lower values for process rates in soil and vegetation than the `intermediate' and `wet' treatments. Regressions showed that soil C-evolution and N-mineralization were positively related to soil moisture content. Not all mineralized N was available for plant uptake in the wet treatment, as a considerable part was denitrified here. Denitrification was especially high as soil water contents increased to levels above field capacity, where redox-potentials sharply dropped. Further, soil P availability was higher under wet conditions. In the `dry' treatment, soil water content was close to the wilting point and plant production was low. In the `intermediate' treatment, plant production was most likely limited by nitrogen. The `wet' treatment did not result in a further increase in plant production. Dam construction and river bed degradation can result in lower river levels and summer drought on floodplains. This experimental study suggests that summer drought on floodplain soils reduces decomposition of soil organic matter, nutrient availability, denitrification, plant production and nutrient uptake. This can affect the capacity of floodplains to remove or retain nutrients from river water in a negative way.     

c-AMP and Morphogenesis of Acetabularia

The c-AMP content has been found to double when Acetabularia develop from 5–10 mm long to grown or almost full-grown algae.
The biological significance of this fact has been approached by studying the effects of drugs known to influence the intracellular c-AMP content on the development of Acetabularia. When grown in the presence of theophyllin or papaverin, inhibitors of phosphodiesterase, the Acetabularia display a striking response during the exponential growth period; the final length, however, is not affected. Both substances increase the c-AMP content of the algea. Isoproterenol, which activates adenylate cyclase in many systems, also influences Acetabularia during the exponential growth period and, in addition, slightly affects cap formation.
The change in c-AMP content in the course of development and the effects of drugs influencing (theophyllin and papaverin) or likely to influence (isoproterenol) the c-AMP content of the algae suggest that this nucleotide plays a role at the time of intense growth.
The same phosphodiesterase activity has been found in the 5–10 mm and the 19–25 mm long algae, whereas two enzymes were found in cap-bearing Acetabularia.
The results are discussed as well as the involvement of c-AMP in the development of this alga.     

Experimental manipulation of water and nutrient input to a Norway spruce plantation at Klosterhede,Denmark

Water and nutrient supply to forest trees are major factors controlling tree growth and forest vitality. Therefore, changes in the supply of water and nutrients to the trees may be important contributing reasons to the forest damages observed in Europe. Such changes may be caused by several factors, e.g. air pollution, soil acidification and climate change. The present study investigates possible effects on tree growth, growth related parameters and nutrition related to changes in the water and nutrient supply. Water and nutrient supply to the forest soil was manipulated in three roof covered plots. The treatments consisted of 1) summer drought, 2) irrigation and 3) combined irrigation and fertilizer application (fertigation). The results from the roof covered treatment plots were compared to a control plot without roof. Increased supply of water during the spring and early summer increased the diameter growth, whereas application of nutrients in addition to irrigation had no additional effect on tree growth. Addition of nutrients increased the needle content of P, Mn, Ca and Mg. Extended summer drought for 2 months had no effect on the tree growth or other growth parameters, but drought reduced the root development in the upper soil layer. Furthermore, drought induced significant stress symptoms by increasing the cone shredding substantially.     

Does sediment resuspension have persistent effects on phytoplankton? Experimental studies in three shallow lakes

1. The persistence of effects of sediment resuspension on chlorophyll a , phytoplankton production and plant nutrients was examined by artificially resuspending sediment in enclosures in three shallow lakes and monitoring concentrations for 4–8 days. Realism was assessed by relating initial suspended solids concentrations to those observed during natural wind events.
2. Positive effects on the phytoplankton, persisting for at least 4–6 days were detected in eight of the nine experiments, and similar effects on nutrient concentrations were also common, although suspended solids had normally returned to control levels within 24 h
3. The phytoplankton during the periods of persistence was normally dominated by planktonic rather than benthic/meroplanktonic genera.
4. Several of the positive responses appeared to be associated with relief of nitrogen deficiency in the algae.
5. Persistent effects from previous resuspension events may cause baseline concentrations of phytoplankton and nutrients in shallow lakes to be overestimated and the effects of resuspension on phytoplankton and nutrients to be underestimated     

Effects of Phosphorus Nutrient on the Hydraulic Conductivity of Sorghum (Sorghum vulgare Pers.) Seedling Roots Under Water Deficiency

Hydroponic experiments were conducted in a growth chamber and changes in the hydraulic conductivity of sorghum (Sorghum vulgare Pers.) roots (Lpr) at the three-leaf stage were measured using the pressure chamber method. Water deficiency was imposed with polyethylene glycol (PEG) 6000 and the phosphorus (P) levels were controlled by complete Hoagland solution with and without P nutrient. The objective of this study was to investigate the effect of P nutrition on root Lpr under water deficiency. The results showed that the Lpr in P deficiency treatments decreased markedly, but the Lpr recovered to the same value as that of control when sufficient P was supplied for 4-24 h. Water deficiency decreased Lpr, but the hydraulic conductivity of the roots with sufficient P supply was still higher than that of plants without P supply. When resuming water supply, the Lpr of the water-deficient plants under P supply recovered faster than that of plants without P supply, which indicates that plants with sufficient P nutrient are more drought tolerant and have a greater ability to recover after drought. The treatment of HgCl2 indicated that P nutrient could regulate the Lpr by affecting the activity and the expression levels of aquaporins.     

Letters to the editor

A decrease in salinity and temperature over the past 3000 years has presented the marine algae of the Baltic Sea with very considerable problems in adaptation. The effects of salinity upon a number of Baltic algae have been measured. The results showed cell mortality to be severe in 0, 68 and 102‰, and minimal in 6 and 11‰: there was most variation in tolerance to 34 and 51‰. The salt tolerances of Baltic marine algae have proved more hyposaline than those of British intertidal algae. Water uptake and loss in tissues of Chorda filum and Fucus vesiculosus from Baltic and British populations have been measured in response to salinity changes. The results revealed significant population differences in both live and killed tissues. Receptacle development and oogonial maturation have been observed in Baltic and British F. vesiculosus, and found to differ in seasonality. Some observations were associated with local sea temperatures but differences in the timing of receptacle initiation and in oogonial size were not. Th depauperate thallus, commonly ascribed to the effects of low salinity, was found to be a complicated phenomenon, comprising numerous attributes which are combined differently in different taxa. The morphological differences between Baltic and British marine algae were usually striking.

The marine algae of the Baltic Sea have therefore diverged in a number of ways from their N. Atlantic counterparts. The naturally high variability of these taxa has enabled them to survive the period of increasingly strong selection pressure which followed the Littorina Sea episode. Divergence seems not to have advanced to the point where speciation may be said to have occurred. The Baltic may therefore be contrasted with the much older Mediterranean Sea, which contains a large number of endemic species. Nevertheless, the Baltic is a site of very considerable evolutionary importance.     

Water translocation in Kalanchoë daigremontiana during periods of drought

Abstract. Kalanchoë daigremontiana strongly reduced daily water loss within 6 d of drought using CAM to restrict transpiration and net CO₂ uptake to the dark period.
Water translocation from old to young leaves of the plant was an additional mechanism which reduced the negative effects of drought on the water relations of young leaves. Excision of old leaves after 7–9 d of drought resulted in a decrease in the water content of young leaves. This was observed despite a decrease in transpirational water loss from young leaves. Water content in young leaves increased slightly in plants with all their leaves in place.
The dry weight of young leaves clearly increased during the experimental period when old leaves were present, but it remained relatively constant in plants without old leaves. Obviously, in addition to water, solutes were transported from old to young leaves of the plant via the phloem. Xylem tension was higher in young compared to old leaves; thus, water translocation could have occurred via xylem elements.
Since transport of organic matter in the phloem is also linked to water flow, phloem transport additionally may contribute effectively to the balance of the water budget in young leaves.