Response of pelagic cyanobacteria to iron additions--enclosure experiments from Lake Erken

In this study, low epilimnetic iron availability during thesummer stagnation period was concluded to be a limiting factorfor cyanobacterial development in Lake Erken. In three enclosureexperiments, different combinations of phosphate, nitrate andammonium additions were tested both with and without additionsof dissolved iron. The addition of iron increased the growthof diazotrophic cyanobacteria significantly compared with enclosuresnot receiving iron. This was especially evident for the colony-formingcyanobacterium Gloeotrichia echinulata. In one experiment, coloniesof G.echinulata disappeared in enclosures not receiving iron,while the abundance of this species increased in those enclosuresto which iron was added.     

RECRUITMENT AND PELAGIC GROWTH OF GLOEOTRICHIA ECHINULATA (CYANOPHYCEAE) IN LAKE ERKEN1

Different parameters in the life cycle of the colony forming cyanobacterium Gloeotrichia echinulata (J.E. Smith) Richter was evaluated in Lake Erken, Sweden. Recruitment of colonies from the sediments and pelagic abundance were measured during 2 years. These data were then used in a model to evaluate and estimate parameters of the life cycle. In our study, recruitment alone only contributed to a small part (<5%) of the maximum G. echinulata abundance that occurred during late summer. However, recruitment from shallow sediments forms the important seed for the pelagic population. Together with measured rates of migration from the sediment, variations in either pelagic colony division rate or pelagic residence time could explain variations in the measured abundance of G. echinulata in situ.     

Temporal changes in biomass specific photosynthesis during the summer: regulation by environmental factors and the importance of phytoplankton succession

Measurements of phytoplankton photosynthesis vs. irradiance relationships have been made at 3–7 day intervals in Lake Erken (central Sweden) for three years during summer stratification. Both the rate of light-limited (^B) and light-saturated (P_max ^B) photosynthesis per unit chlorophyll a showed distinct and similar temporal trends in each year. Seasonal trends were especially evident for P_max ^B, which increased in value for several weeks following the onset of thermal stratification, and then declined in the presence of the large colonial blue-green alga, Gloeotrichia echinulata. By late summer, when the biomass of G. echinulata had decreased, P_max ^B again rose to its early summer value. The covariation of biomass-specific photosynthesis with the blooming of G. echinulata was the one clear seasonal (week-month) pattern which emerged in each of 3 years. Over short (day-week) time scales, changes in ^B were related to changes in irradiance exposure on the day of sampling. However, the relationship between these two parameters was variable in time, since it was superimposed upon longer term trends controlled by changes in phytoplankton species composition. Increases in G. echinulata biomass corresponded with a deepening of the thermocline, which both increased internal phosphorus loading and the transport of resting G. echinulata colonies into the epilimnion. The timing and magnitude of the yearly G. echinulata bloom was as a result related to the seasonal development of thermal stratification. These results illustrate the importance of seasonal changes in the phytoplankton community as a factor regulating rates of biomass specific photosynthesis, particularly when the successional changes involve species with very different life strategies.     

Gloeotrichia echinulata, a colonial cyanobacterium with a unique phosphorus uptake and life strategy

Epilimnetic colonies of Gloeotrichia echinulata were harvestedfrom 200–300 I of water in Lake Erken with filtrationthrough appropriate plankton nets (200 or 70 µm). Phosphateuptake characteristics, phosphorus (P) status and photosynthesisof the colonies were determined twice a week during July andAugust 1991. Phosphate uptake was analysed according to thesimple force- flow relationship of Falkner et al. (Arch. Microbiol.,152, 353–361, 1989). The threshold concentration of Puptake below which uptake ceases for energetic reasons, wasconstantly much higher than the epilimnetic soluble reactivephosphorus (SRP) concentration, so that the planktonic colonieswere unable to acquire any Pin the epilimnion. Neither did organicP seem to be a source of P for planktonic colonies. Gloeotrichiaechinulata has a unique life strategy in comparison to othercommon genera of bloom-forming cynanobacteria. Its P assimilationand growth are completely separated both in time and space;growth is preceded by benthic P assimilation. Epilimnetic growthwas based solely on internally stored P and growth rates fittedthe Droop model well. Depletion of stored P restricts the lengthof the planktonic phase to 15–20 days under ‘optimal’growth conditions. Wind-induced surface drift seemed to be themost important loss factor from the epilimnion. Massive recruitmentof P-rich benthic colonies accounted for two-thirds of totalnet internal P loading observed between mid-July and mid-August(3.8 mg P m^–2 day^–1).     

羊草绿叶氮、磷浓度和比叶面积沿氮、磷和水分梯度的变化

全球气候变化包括氮沉降增加和降水格局改变, 影响着生态系统的生物地球化学循环, 进而可能会对植物的养分保持特性产生影响。研究不同氮、磷和水分梯度上植物叶片养分特性的响应格局, 对于预测氮沉降增加和降水格局的改变对植物养分策略的潜在影响, 具有一定的理论与实践意义。该研究通过3个盆栽控制实验 (施氮肥实验:0、0.5、1、2、4、8、16、32、64和128g·m-2等10个水平;施磷肥实验:梯度同施氮肥实验;控水实验:3600、 4000、4500、5143、6000、7200、9000、12000、18000和36000ml-pot-1等10个水平), 探讨了羊草 (Leymuschinensis) 绿叶氮、磷浓度和比叶面积 (Specificleafarea, SLA) 对这些环境因子改变的响应。结果表明, 在一定范围内, 施氮肥提高了羊草绿叶比叶面积和绿叶氮浓度, 但对绿叶磷浓度没有明显的影响;施磷肥显著提高了绿叶比叶面积、绿叶氮浓度和绿叶磷浓度;供水量增加提高了羊草绿叶比叶面积, 降低了绿叶氮浓度, 但对绿叶磷浓度无显著影响。这表明, 氮、磷和水分因子的改变影响了植物叶片的养分保持能力, 且不同梯度的影响程度也不同。因此, 全球气候变化可能影响植物养分利用策略, 进而可能对植被-土壤系统养分循环产生影响。     

Response of the cyanobacterium Gloeotrichia echinulata to iron and boron additions – an experiment from Lake Erken

1. This study considers whether the availability of iron and boron are important influences on the development of the cyanobacterium Gloeotrichia echinulata in Lake Erken, Sweden.
2. In an in situ experiment, phosphate and nitrate were added to all enclosures, but pelagic colonies of G. echinulata only increased in abundance in enclosures to which iron had also been added.
3. An even greater increase in the abundance of G. echinulata occured in enclosures in which the additions of phosphate, nitrate and iron were complemented by additions of boron.
4. Boron additions, together with phosphate and nitrate but without iron, did not stimulate the growth of G. echinulata .     

The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

Factors controlling pelagic populations of ciliates and heliozoans-- late summer investigations in an acidic lake before and after liming

Ciliates and heliozoans were studied in a small oligotrophicSwedish lake with the objective of investigating what controlledthe community structure and population dynamics. Enclosure experimentswere performed with five levels of inorganic nutrient concentrationsand the presence or absence of crustacean zooplankton. The experimentswere performed during the summers immediately preceding andfollowing liming of the lake. The responses of protozooplanktonto these manipulations were investigated and compared with theresponses of bacterioplankton, phytoplankton and metazooplankton.In contrast to most studies on protozoan zooplankton, ciliateswere determined to the lowest taxon possible, and biomassesand abundances were calculated for the different taxa. The protozooplanktonbiomass of the lake, expressed as summer average, did not changebetween the years. However, the community composition changedfrom smaller to larger species as the small prostomatid ciliatesUrotricha and Balanion were largely replaced by oligotrichsof the genera Strombidium and Strobilidium. The experimentssuggested that the protozooplankton community in the lake wascontrolled by metazooplankton in the acidic environment, whilefood limitation was the most important controlling factor afterliming. Before liming, Urotricha and Balanion were unaffectedby nutrient additions as well as by metazooplankton, while otherciliates and Heliozoa showed a significantly negative responseto metazooplankton. After liming, Urotricha and Balanion showeda significant positive response to nutrient addition as wellas to bacterioplankton biomass. A possible explanation for thisresponse in these algivore genera is that the response was indirectand mediated through small chrysoflagellates.     

Sediment-to-water blue-green algal recruitment in response to alum and environmental factors

The sediment-to-water recruitment of blue-green algae was investigated in a shallow lake following treatment with aluminum sulfate and sodium aluminate to control sediment phosphorus (P) release. A comparison of results from two summers each before and after treatment indicates that the treatment did not universally impact the recruitment of either sporulating or non-sporulating forms of blue-green algae. Blooms of Anabaena, Aphanizomenon, and Coelosphaerium resulted predominantly from growth in the water column following strong recruitment episodes lasting up to two weeks, while Microcystis populations were relatively insensitive to periodically high inputs from recruitment. The development of planktonic populations of Gloeotrichia echinulata, by contrast, were largely dependent on sustained recruitment in response to adequate light and temperature regimes at the sediment surface.The cellular P content of recruited G. echinulata colonies was unaffected by the accumulation of aluminum floc to the lake sediments. Both G. echinulata and C. naegelianum showed elevated levels of cellular P in newly recruited colonies as compared to planktonic colonies, indicating P transport from the sediments to the water column. Total P translocation by blue-green algae was negligible in the absence of a substantial recruitment of G. echinulata. The recruitment of G. echinulata, and hence the magnitude of P translocation, was therefore more responsive to environmental conditions prevalent at the sediments than to direct effects of the treatment itself.     

The role of Gloeotrichia echinulata in the transfer of phosphorus from sediments to water in Lake Erken

The abundance of Gloeotrichia echinulata colonies in the sediments of Lake Erken and their phosphorus content were investigated to determine the contribution of Gloeotrichia colonies to total sediment phosphorus. Moreover, the potential size of the algal inoculum and the migration to the water during summer were estimated.The surplus phosphorus content of the resting colonies in the sediment was about 45% of total phosphorus, which maximized at 8.5 µg P (mg dw)^–1 or 81 ng P colony^–1. The C:P ratio (by weight) in the early colonies appearing in the lake water was 50:1, while the ratio stabilized at 150 during the major migration period. The internal supply of surplus phosphorus was used during the pelagic growth of the colonies.The internal phosphorus loading to the epilimnion of Lake Erken due to Gloeotrichia migration could, from the measurements of the increase in particulate epilimnetic phosphorus, be estimated at 40 mg P m ^–2 or 2.5 mg P m^–2 d^–1 in late July and early August. Determination of the number of colonies in the sediment before and during the migration verified this value to be a conservative estimate of the internal phosphorus loading due to Gloeotrichia migration to the epilimnion in Lake Erken.The sediment P content calculated from the P concentration in early epilimnion colonies resulted in a value of 35 µg P (g dw)^–1 as a maximum. This corresponds to only 3% of the total phosphorus content in Lake Erken sediment.     

Effects of Desiccation on Phosphorus and Potassium Acquisition by a Desiccation-tolerant Moss and Lichen

The hypothesis that desiccation-tolerant mosses and lichensmay be more responsive to nutrient inputs accompanying intermittentdesiccation than mesophytic forest species was investigatedemploying species from semi-arid grassland in Hungary. Shootapices of the moss Syntrichia ruralis and marginal lobes ofthe lichen Cladonia convoluta were maintained for 7 weeks undercontrolled conditions. They were cultivated with or withouta weekly application of the major inorganic macronutrients,and either under constant hydration or with one or two 24 hperiods of desiccation each week. Growth of S. ruralis was stimulatedby nutrient additions, but lower weight increments were achievedwith increasing frequency of desiccation. All samples of thelichen showed positive growth, and no significant treatmenteffects were detected. A large net uptake of P occurred in nutrient-treatedmaterial of both species that was unaffected by the impositionof desiccation treatments. A smaller net uptake of K into theintracellular fraction was also observed when nutrients wereapplied, but in the moss this was against a baseline of decreasingK content. In contrast, more of the original K content was retainedin C. convoluta. In neither species was any clear evidence foundfor inhibition of nutrient uptake by the desiccation episodes.It is suggested that the lack of growth response in the lichenarises from an inability to bring together the additional nutrients,presumably mainly absorbed by the mycobiont, with photosynthateproduced by the photobiont. Copyright 2000 Annals of BotanyCompany Cladonia convoluta, Syntrichia ruralis, desiccation tolerance, mineral nutrition, phosphorus, potassium     

Diurnal variations in the auto- and heterotrophic activity of cyanobacterial phycospheres (Gloeotrichia echinulata) and the identity of attached bacteria

1. We assessed the role of cyanobacterial–bacterial consortia (Gloeotrichia echinulata phycospheres) for net changes in inorganic carbon, primary production (PP) and secondary production in Lake Erken (Sweden). 2. At the time of sampling, large colonies of G. echinulata formed a massive bloom with abundances ranging from 102 colonies L^?1 in the pelagic zone to 5000 colonies L^?1 in shallow bays. These colonies and their surrounding phycospheres contributed between 17 and 92% of total PP, and phycosphere‐associated bacteria contributed between 8.5 and 82% of total bacterial secondary production. PP followed a diurnal cycle, whereas bacterial production showed no such pattern. Over a 24 h period, carbon dioxide measurements showed that the phycospheres were net autotrophic in the top layer of the water column, whereas they were net heterotrophic below 2 m depth. 3. Sequencing and phylogenetic analysis of 16S rRNA genes of attached bacteria revealed a diverse bacterial community that included populations affiliated with Proteobacteria, Bacteriodetes, Acidobacteria, Fusobacteria, Firmicutes, Verrucomicrobia, and other Cyanobacteria. 4. Compared with their planktonic counterparts, bacteria associated with cyanobacterial phycospheres had lower affinity for arginine, used as a model compound to assess uptake of organic compounds. 5. Extrapolation of our data to the water column of lake Erken suggests that microorganisms that were not associated with cyanobacteria dominated CO₂ production at the ecosystem scale during our experiments, as CO₂ fixation balanced CO₂ production in the cyanobacterial phycospheres.     

Nutrient Dilution by Starch in CO2-enriched Chrysanthemum

Increasing growth irradiance and CO₂ generally decreases foliarnutrient concentration on a dry weight basis and increases foliarstarch concentration. However, the extent to which starch concentrations‘dilute’ foliar nutrient concentrations when thelatter are expressed on a dry weight basis is not known. Todetermine the importance of differential starch accumulationin calculating nutrient concentrations on a dry weight basis,leaf nutrient and starch concentrations were measured in Chrysanthemum? morifolium ‘Fiesta’ (Ramat.) cuttings grown atthree irradiance levels and two CO₂ levels for eight weeks inboth winter and spring. On a dry weight basis, foliar concentrationsof most nutrients were lower in both seasons as a result ofthe elevated CO₂ and irradiance levels, and total dry weightswere higher. Per cent starch was greater at the high CO₂ levelin both seasons but was only greater at higher irradiances inthe winter experiment. When starch was subtracted from the leafdry weights, the differences between CO₂ and irradiance treatmentsdisappeared with respect to N, P, K, Ca, Mg, S, and B but notfor Fe, Mn, Zn, and Cu. Key words: CO₂ enrichment, starch, nutrients, irradiance     

Importance of glucose-6-phosphate dehydrogenase activity in cell death

The intracellular redox potential plays an important role incell survival. The principal intracellular reductant NADPH is mainlyproduced by the pentose phosphate pathway by glucose-6-phosphate dehydrogenase (G6PDH), the rate-limiting enzyme, and by6-phosphogluconate dehydrogenase. Considering the importance of NADPH,we hypothesized that G6PDH plays a critical role in cell death. Ourresults show that 1) G6PDHinhibitors potentiatedH₂O₂-inducedcell death; 2) overexpression ofG6PDH increased resistance toH₂O₂-induced cell death; 3) serum deprivation, astimulator of cell death, was associated with decreased G6PDH activityand resulted in elevated reactive oxygen species (ROS);4) additions of substrates for G6PDHto serum-deprived cells almost completely abrogated the serumdeprivation-induced rise in ROS; 5)consequences of G6PDH inhibition included a significant increase inapoptosis, loss of protein thiols, and degradation of G6PDH; and6) G6PDH inhibition caused changesin mitogen-activated protein kinase phosphorylation that were similarto the changes seen withH₂O₂.We conclude that G6PDH plays a critical role in cell death by affectingthe redox potential.     

Ecological investigations of blooms of colonial Phaeocystis pouchetti--I. Abundance, biochemical composition, and metabolic rates

A winter bloom of the colonial stage of the prymnesiophyte Phaeocystispouchetit was studied in the 13-m³ mesocosms of the Marine EcosystemResearch Laboratory on Narragansett Bay, Rhode Island The tankswere temperature regulated at 4±2°C but differedin their nutrient concentrations and in situ irradiances. Oneof the tanks was a control without added nutrients, one receiveda temporary nutrient spike and two others received daily N/P/Siinputs. Photosynthesis and growth rates of colonies exposedto a range of natural light levels were measured at weekly intervals.Particulate carbon production and release of dissolved organiccarbon (DOC) by the entire plankton community was determinedconcurrently. Photosynthesis and growth rates of Phaeocystisin tanks receiving daily nutrient additions were asymptoticfunctions of irradiance. Light-saturated rates exhibited asymptoticrelationships with dissolved inorganic nitrogen (N) levels.N-Limited populations showed more variable responses. Althoughirradiance and N availability regulated the population dynamicsof Phaeocystis, the presence or absence of silicate (S₁) influencedits relative importance in each tank. Phaeocystis dominatedcommunity metabolism in the absence of Si, but co-occurred withextensive stands of diatoms when Si was available. A significantpositive correlation was found between the contribution by Phaeocystisto community production and the proportion of photosynthatereleased as DOC In all tanks, Phaeocystis populations exhibitedcycles of abundance in which division of cells within coloniespreceded the multiplication of colonies. The production of newcolonies apparently occurred via two mechanisms: the formationof colonies from solitary cells, and the cleavage of largercolonies into smaller daughter colonies. Phaeocystis in tankswith near undetectable nutrient levels contained C:N, C:Chla, and C:ATP ratios several times higher than colonies in nutnent-repletetanks. Phaeocystis C:Chl a and C:ATP ratios were substantiallygreater than those of non-gelatinous phytoplankton due to carbohydratestorage in colony gelatin In contrast, C:N ratios in Phaeocystisand non-gelatinous phytoplankton were similar, suggesting astorage depot of organic N outside of the cells. The resultssupport the notion that Phaeocystis colonies function as biologicalentities rather than as passive aggregations of cells.     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.     

Role of the pentose phosphate pathway during callus development in explants from potato tuber

Metabolic change during callus formation in explants from potatotuber (Solarium tuberosum L.) was investigated. Under the presentculture conditions, callus starts exponential growth (estimatedby the fresh weight increase) with a 5-day lag period, at whichtime marked production of DNA occurs. In the lag period, respirationis very much resistant to malonate, and the value of the C6/C1ratio and activities of both G6PDH 6PGDH pass their peaks. Onthe basis of these and relevant findings, glucose metabolismthrough the PP pathway is surmised to play a significant rolein the initial DNA multiplication phase of callus development. (Received October 15, 1969; )     

Nutrient and phytoplankton dynamics in the Queen Charlotte Islands (Canada) during the summer upwelling seasons of 2001-2002

The Queen Charlotte Islands, Canada, lie at the northern extremeof the coastal upwelling system of the eastern North PacificOcean (51–54°N). In this study, the first observationsof spatial and inter-annual patterns in nutrient inventories,chlorophyll (Chl) a and phytoplankton assemblages are reportedand related to oceanographic conditions in near-shore watersof the island archipelago. Stronger and more persistent upwellingin 2002 coincided with higher nutrient and Chl a standing stockscompared to 2001 and a higher proportion of diatoms. Dinoflagellateswere more prevalent in 2001, including several potentially harmfulspecies. At sub-seasonal scales, Chl a concentrations were greatestduring downwelling conditions and smallest during upwellingconditions. On the west coast, weak water column stratification,high relative proportions of diatoms and large nutrient inventoriesaccompanied upwelling-favourable conditions, whereas on theeast coast, there was no direct relationship between the BakunUpwelling Index and water column stability. According to redundancyanalysis, variability in species composition was best explainedby sea surface temperature, the depth of the euphotic zone andnutrient inventories. The east coast supported blooms of coccolithophoreswithin protected bays, confirming previous satellite observationsthat showed bright patches in these areas. The data illustratethat moderate upwelling can have an important influence on near-shorealgal standing stocks and species composition at the northernextreme of the upwelling system off the west coast of NorthAmerica, and that topographical complexity may be importantfor the development of phytoplankton blooms.     

Factors regulating recruitment from the sediment to the water column in the bloom-forming cyanobacterium Gloeotrichia echinulata

1. The influence of light, temperature, sediment mixing and sediment origin (water depth) on the recruitment of the cyanobacterium Gloeotrichia echinulata was examined in the laboratory. 2. Light and temperature were the most important factors initiating germination in G. echinulata. 3. The extent of germination (recruited biovolume) was mainly regulated by temperature and sediment mixing. Furthermore, sediment mixing significantly enhanced the frequency of observed heterocysts and colonies. 4. Despite the fact that the deep and shallow sediments contained a similar number of akinete colonies, the highest recruitment occurred from shallow sediments, indicating that akinetes from shallow sediments have a higher viability than those from deeper parts of the lake. 5. Our results support the hypothesis that shallow sediments are more important than profundal sediments for the recruitment of G. echinulata to the pelagic zone.     

The spatial and temporal dynamics of deep chlorophyll layers in high-mountain lakes: effects of nutrients, grazing and herbivore nutrient recycling as growth determinants

Deep chlorophyll layers (DCL) are a common feature of oligotrophiclakes, yet the mechanisms that form and maintain them are notunderstood fully. These phytoplankton populations occur in themetalimnia of lakes where light levels are moderate to low,and where nutrient levels and zooplankton grazing pressure aredifferent than in the epilimnion. To test the importance ofnutrients and grazing pressure for algal growth in differentlake strata, microcosm experiments and monitoring were conductedin two oligotrophic lakes in the Rocky Mountains of North Americathat contain DCL. In situ microcosm experiments with naturalphytoplankton communities from three depth strata were conductedwith macronutrient additions and with and without the naturalzooplankton grazing communities. Alkaline phosphatase assaysand the in situ microcosm experiments indicated less nutrientlimitation in the metalimnia than in the epilimnia of both lakes.Zooplankton grazing in the experiments decreased algal populationgrowth rates by as much as 6% day^–1, with impacts shiftingto progressively deeper strata over the summer. Zooplanktongrazing losses, however, were partially offset by nutrient recyclingthat increased algal growth rates. Depth-differential nutrientdeficiency and zooplankton grazing and recycling interactedto maintain the DCL in these lakes.