The effects of catchment liming on the chemistry and biology of upland Welsh streams: testing model predictions

1. The catchments of three acidified streams in mid Wales were limed in 1987/88. Here we assess their chemical and biological response in comparison with unmanipulated reference streams over a period of 5 years post-liming. 2. Stream chemistry was measured weekly/fortnightly between 1985 and 1992, while macroinvertebrates were sampled annually. Colonization by acid-sensitive taxa was assessed and trends in community structure were monitored by TWINSPAN. Real biological responses were compared with those predicted by an empirical model constructed using chemical data. 3. There were marked changes in stream chemistry following liming: calcium concentrations and pH values increased, while aluminium concentrations decreased to levels similar to those in naturally circumneutral streams. These conditions have persisted since liming. 4. Empirical models predicted that stream invertebrates would respond to the altered stream chemistry, with the establishment of communities typical of circumneutral conditions. 5. Following liming, the taxon richness and abundance of acid-sensitive taxa was significantly higher in limed compared with reference streams. Colonization by, and persistence of acid-sensitive taxa was patchy, however, and richness was still substantially lower than in naturally circumneutral streams. Moreover, contrary to the model predictions, there were no wholesale changes in the structure of macroinvertebrate communities. 6. We conclude that liming has created and maintained chemical conditions suitable for macroinvertebrate communities typical of circumneutral streams, but these chemical changes have not been matched by sustained responses among the biota. The views expressed in this article are those of the authors and do not necessarily reflect those of the NRA.     

The effects of low pH and palliative liming on beech litter decomposition in acid-sensitive streams

The decomposition of allochthonous leaf litter is retarded by stream acidification, but few studies have evaluated whether this effect can be offset by liming – the palliative addition of calcium carbonate either to streams or their catchments. We assessed the response of litter decomposition to pH and experimental liming in Welsh upland streams. Small-mesh (<335 μm) litter-bags containing common beech (Fagus sylvatica L.) were submerged in main river sites along the River Wye, and in replicate acid, circumneutral and experimentally limed tributaries (all n = 3) for 20 days. Beech decomposition was inhibited in acid tributaries and main river sites compared to circumneutral tributaries. Despite having only moderately increased pH relative to acid streams, limed sites had increased decomposition rates that were indistinguishable from naturally circumneutral streams. Decomposition rates increased highly significantly with pH across all 12 sites studied, and values were near identical to those in more prolonged experiments elsewhere. There were no significant variations in shredder numbers with decomposition rate, and no evidence that sites with faster decomposition had smaller shredder proportions. Although based on short-term observations and leaves from just one tree species, these results are consistent with the well-known retardation at low pH of some aspect microbial decomposition (e.g. by hyphomycete fungi). They are among the first to suggest that stream liming to combat acidification might reverse such impacts of low pH. Further data are required on the microbiological causes and ecological consequences of altered detrital processing in acid-sensitive and limed streams.     

Acute effects on perch (Perca fluviatilis) and long-term effects on whitefish (Coregonus lavaretus pallasi) of liming of an acidified lake

The acute effects on perch and long‐term effects on whitefish were studied after liming one side of a lake [initial pH: 4.6–5.5; aluminium (Al): 29–54 μg L^?1; Ca²⁺: 0.02–0.07 mmol L^?1] that was divided into two parts with a plastic curtain. Some Al precipitation was observed on the gill surface of perch 1 day after liming; the concentration of plasma sodium of perch females in the limed side was also higher than in the acidic side. Nearly 6 months after liming, at the spawning time of whitefish, the plasma chloride concentration of whitefish in the limed side was higher and blood glucose concentration lower than in the acidic side. Four of five whitefish males from the limed side, but only one of five males and none of the five females from the acidic side, were ready to spawn. The growth of whitefish in the limed side was also more rapid. These changes illustrate that liming had no acute harmful effects on perch, and allowed whitefish to recover from acidity‐related physiological stress.     

Limestone Treatment of Whetstone Brook, Massachusetts. I. Treatment Methodology and Water-Chemistry Changes during Treatment

Whetstone Brook is a trout stream located in north-central Massachusetts that is degraded by acid precipitation. The stream was treated with 56 tonnes of powdered limestone by a prototype, water-powered doser as part of the U.S. Fish and Wildlife Service's Acid Precipitation Mitigation Program. The goal of liming Whetstone Brook was to raise the pH to 6.5 and acid neutralizing capacity (ANC) to at least 50 μeq/1 in a 3.2-km reach. This goal was achieved despite the fact that during the 31 months of treatment stream flow was 37% higher than during the pretreatment period. During the treatment period, pH averaged 6.54 and ANC averaged 69.75. During the pretreatment period average pH was 5.97 and average ANC was 20.26. In the control section of Whetstone Brook, both pH and ANC were lower during the treatment period than during the pretreatment period. During treatment, monomeric aluminum, a form toxic to fish, declined in the treated section and increased in the control section. Total calcium, sediment calcium, and pore-water calcium increased in the treated section during treatment but declined in the control section. The other base anions and cations, nutrients, and physical parameters were not significantly affected by liming.     

Changes in stream chemistry and biology in response to reduced levels of acid deposition during 1987–2003 in the Neversink River Basin,Catskill Mountains

Atmospheric acid deposition has decreased in the northeastern United States since the 1970s, resulting in modest increases in pH, acid-neutralizing capacity (ANC), and decreases in inorganic monomeric aluminum (Al_IM) concentrations since stream chemistry monitoring began in the 1980s in the acid-sensitive upper Neversink River basin in the Catskill Mountains of New York. Stream pH has increased by 0.01 units/year during 1987–2003 at three sites in the Neversink basin as determined by Seasonal Kendall trend analysis. In light of this observed decrease in stream acidity, we sampled 12 stream sites within the Neversink River watershed for water chemistry, macroinvertebrates, fish, and periphytic diatoms in 2003 to compare with a similar data set collected in 1987. Metrics and indices that reflect sensitivity to stream acidity were developed with these biological data to determine whether changes in stream biota over the intervening 16 years parallel those of stream chemistry. Statistical comparisons of data on stream chemistry and an acid biological assessment profile (Acid BAP) derived from invertebrate data showed no significant differences between the two years. For pH and ANC, however, values in 2003 were generally lower than those in 1987; this difference likely resulted from higher streamflow in summer 2003. Despite these likely flow-induced changes in summer 2003, an ordination and cluster analysis of macroinvertebrate taxa based on the Acid BAP indicated that the most acidic sites in the upstream half of the East Branch Neversink River form a statistically significant separate cluster consistent with less acidic stream conditions. This analysis is consistent with limited recovery of invertebrate species in the most acidic reaches of the river, but will require additional improvement in stream chemistry before a stronger conclusion can be drawn. Data on the fish and periphytic diatom communities in 2003 indicate that slimy sculpin had not extended their habitat to upstream reaches that previously were devoid of this acid-intolerant species in 1987; a diatom acid-tolerance index indicates continued high-acid impact throughout most of the East Branch and headwaters of the West Branch Neversink River.     

Assessment of mine drainage remediated streams using diatom assemblages and biofilm enzyme activities

A legacy of pre-regulation coal mining in many areas has been acid mine drainage (metal-rich, low pH water). Today, numerous remediation strategies may be implemented, but there is little data on efficacy in restoring biological condition. Two alkaline doser projects in Ohio were assessed using diatom assemblages, and biofilm extracellular enzyme activities (EEA). In one stream, water quality steadily increased downstream of the doser; pH increased from 3.8 to 7.2 and Fe decreased from 107 to 0.42 mg l^?1. Likewise, the periphyton biomass (chl a 7.15–12.77 mg m^?2) increased and periphyton index scores (4–27) improved. As well, EEA data showed greater activity for phosphorus, nitrogen, and one carbon acquisition enzyme. For the other stream, the conductivity remained high (>720 μS cm^?1) and pH and alkalinity decreased downstream. Biological data, including EEA, varied along the stream length with higher numbers in the middle reach, such as chlorophyll a (0.56 to 87.75 to 2.77 mg m^?2), and index scores (7 to 29 to 11). The first remediation site showed positive results in chemistry, biological community and measures of ecosystem function. The second stream was highly variable in these parameters suggesting further AMD inputs are hampering recovery.     

Long-term trends in limnological characteristics in the Aurora trout lakes, Sudbury, Canada

Paleolimnological techniques were employed to document the limnological histories of the aurora trout lakes, located in the Sudbury region of Ontario. Two of these lakes are of special interest to fisheries managers, as they represent the only known native habitats of a rare strain of brook trout: the aurora trout. These lakes were limed as part of restoration efforts. Stratigraphic changes in diatom and chrysophyte assemblages from dated lake sediment cores indicate that all the lakes have been impacted by anthropogenic acidification, although the timing and the magnitude of acidification were different amongst the lakes. For example, Whirligig Lake was likely the most naturally acidic lake in the past, but it had further acidified since about 1960. This lake was limed in 1989 and then again in 1993. In Whitepine Lake, acidification started 1940; however, in the most recent sediments (1992), some recovery in lakewater acidity has occurred. In Little Whitepine Lake (a reference lake), acidification started earlier (1920) and the lakewater pH continued to decline until about 1990. This lake was limed in 1989. The chrysophyte paleoindicators suggest a recent recovery in this lake. The successful re-introduction of aurora trout in Whirligig and Whitepine lakes is undoubtedly related to the improved water quality through liming but, based on our paleolimnological indicators, the lakes' limnological characteristics (e.g. pH and metal concentrations) are still different from those present before atmospheric deposition of strong acids from the Sudbury smelters.     

Long-term effects of catchment liming on invertebrates in upland streams

1. Catchment liming to mitigate acidification causes major chemical change in freshwaters but longer‐term effects are poorly understood. Using a replicated basin‐scale experiment with a multiple BACI design (= before‐after‐control‐impact), we assessed chemical and biological effects for 10 years after the catchments of three acidified Welsh streams at Llyn Brianne were limed in 1987/88. 2. Stream chemistry was measured weekly to monthly, and macroinvertebrates monitored annually, between 1985 and 1998. Biological change through time was assessed from the abundance and taxon richness of invertebrates. We paid particular attention to 18 species known to be acid‐sensitive. The effects of liming were assessed by comparing chemical and biological trends among the three replicate limed streams, three acid reference streams and two naturally circumneutral streams. 3. Following single lime applications, acid‐base chemistry in treated streams changed significantly. High mean pH (> 6), increased calcium (> 2.5 mg L^?1) and low aluminium (< 0.1 mg L^?1) persisted throughout the 10 years following liming. 4. The effects of liming on invertebrates were modest. Acid sensitive taxa increased significantly in abundance in limed streams, but only during 2 years following treatment. Significant effects on richness were more sustained, but on average added only 2–3 acid‐sensitive species to the treated streams, roughly one‐third of their average richness in adjacent circumneutral streams. Only the mayfly Baetis rhodani and the stonefly Brachyptera risi occurred significantly more often in limed streams after treatment than before it. 5. Despite these modest long‐term effects on invertebrates, nearly 80% of the total pool of acid‐sensitive species has occurred at least once in the limed streams in the 10 years since treatment. This pattern of occurrence suggests that the colonization of limed streams by acid‐sensitive taxa reflects limited persistence rather than restricted dispersal. We present evidence to show that episodes of low pH continued to affect acid‐sensitive taxa even after liming. We highlight the importance of extending the time‐periods over which the effects of large‐scale ecological experiments are assessed.     

Dissolved organic carbon, seston and macroinvertebrate drift in an acidified and limed humic stream

SUMMARY. 1. Total seston, and invertebrate drift were studied before and after lime addition to Fyllean River, a stream-iake system in Halland county, southwest Sweden, with poorly buffered waters undergoing acidification. 2. The largest effect of liming was on the chemistry of the water. Following liming with 23 mg CaCO₃ l^?1 the pH of the water changed from 5.8 to 6.8 and alkalinity from 0.04 to 0.13 meq l^?1.Turbidity increased from 3.4 to 4.7 JTU with no change in colour. 3. Dissolved organic carbon (DOC) concentration of all samples was in the range 10.7–13.3 mg C l^?1 with no significant change occurring due to liming. 4. Total seston increased from 4.35 mg DM 1^?1 in unlimed conditions to 6.25 mg DM l^?1 after lime addition. All significant changes in seston occurred in the smaller size fraction (0.45–25 μm). 5. Liming reduced the organic content of the partieulate material from an average of 61% to 39% immediately downstream of a lime silo (within 1 km) but had little effect when the river course was interrupted by lakes and impoundments. 6. The lakes in the river system had a larger effect on seston concentration than any effect of the lime addition by itself. Particle concentrations were reduced by 50–55% and DOC by about 1 mg C l^?1as the water passed through the lakes. 7. Macroinvertebrate drift density was low in all samples before and after liming and typical of oligotrophic streams. Drift was significantly lower at limed (0.024 ind. m^?3) than at unlimed (0.083 ind. m^?3) locations. The decrease was only in total drift density with no significant change in the relative abundance of functional groups or in densities of single taxa, except for a reduction in drift of predators in the limed condition.     

Lake responses to long‐term disturbances and management practices

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Effect of whole catchment liming on the episodic acidification of two adirondack streams

During the fall of 1989 7.7Mg/ha of calcium carbonate was applied on two tributary catchments (40 ha and 60 ha) to Woods Lake, a small (25 ha) acidic headwater lake in the western Adirondack region of New York. Stream-water chemistry in both catchment tributaries responded immediately. Acid-neutralizing capacity (ANC) increased by more than 200 eq/L in one of the streams and more than 1000 eq/L in the other, from pre-liming values which ranged from –25 to +40 eq/L. The increase in ANC was primarily due to increases in dissolved Ca²⁺ concentrations. Most of the initial response of the streams was due to the dissolution of calcite that fell directly into the stream channels and adjacent wetlands. A small beaver impoundment and associated wetlands were probably responsible for the greater response observed in one of the streams.After the liming of subcatchmentIV (60 ha), Ca²⁺ concentrations increased with increasing stream discharge in the stream during fall rain events, suggesting a contribution from calcite dissolved within the soil and transported to the stream by surface runoff or shallow interflow. Concentrations of other ions not associated with the calcite (e.g. Na⁺) decreased during fall rain events, presumably due to mixing of solute-rich base flow with more dilute shallow interflow. The strong relation between changes in Ca²⁺ and changes in NO ₃ ^– concentrations during spring snowmelt, (r² = 0.93, slope = 0.96, on an equivalent basis) suggests that both solutes had a common source in the organic horizon of the soil. Increases in NO ₃ ^– concentrations during snowmelt were balanced by increases in Ca²⁺ that was released either directly from the calcite or from exchange sites, mitigating episodic acidification of the stream. However, high ambient NO ₃ ^– concentrations and relatively low ambient Ca²⁺ concentrations in the stream during the spring caused the stream to become acidic despite the CaCO₃ treatment.In stream WO2 (40ha), Ca²⁺ concentrations were much higher than in stream WO4 because of the dissolution of calcite which fell directly into the upstream beaver pond and its associated wetlands. Calcium concentrations decreased as both NO ₃ ^– concentrations and stream discharge increased, due to the dilution of Ca-enriched beaver pond water by shallow interflow. Despite this dilution, Ca²⁺ concentrations were high enough to more than balance strong acid anion (SO ₄ ^– , NO ₃ ^– , Cl^–) concentrations, resulting in a positive ANC in this stream throughout the year. These data indicate that liming of wetlands and beaver ponds is more effective than whole catchment liming in neutralizing acidic surface waters.     

Increases in soil organic carbon sequestration can reduce the global warming potential of long‐term liming to permanent grassland

The application of calcium‐ and magnesium‐rich materials to soil, known as liming, has long been a foundation of many agro‐ecosystems worldwide because of its role in counteracting soil acidity. Although liming contributes to increased rates of respiration from soil thereby potentially reducing soils ability to act as a CO₂ sink, the long‐term effects of liming on soil organic carbon (C_org) sequestration are largely unknown. Here, using data spanning 129 years of the Park Grass Experiment at Rothamsted (UK), we show net C_org sequestration measured in the 0–23 cm layer at different time intervals since 1876 was 2–20 times greater in limed than in unlimed soils. The main cause of this large C_org accrual was greater biological activity in limed soils, which despite increasing soil respiration rates, led to plant C inputs being processed and incorporated into resistant soil organo‐mineral pools. Limed organo‐mineral soils showed: (1) greater C_org content for similar plant productivity levels (i.e. hay yields); (2) higher ¹⁴C incorporation after 1950s atomic bomb testing and (3) lower C : N ratios than unlimed organo‐mineral soils, which also indicate higher microbial processing of plant C. Our results show that greater C_org sequestration in limed soils strongly reduced the global warming potential of long‐term liming to permanent grassland suggesting the net contribution of agricultural liming to global warming could be lower than previously estimated. Our study demonstrates that liming might prove to be an effective mitigation strategy, especially because liming applications can be associated with a reduced use of nitrogen fertilizer which is a key cause for increased greenhouse gas emissions from agro‐ecosystems.     

Chemical characteristics of lakes in the High Tatra Mountains,Slovakia

The chemistry of 53 lakes at various stages of acidification and inhabited (at the presence and/or in the past) by pelagic Crustacea was studied in September 1984. Ten of these lakes were investigated in detail biannually (July and October 1987–1990). The July results reflect the influence of snowmelt and were compared with the October ones. The most important anion was sulphate with the average values of 98 and 104 μeq 1^-1 in 1984 and 1987–1990, respectively. High concentrations of nitrate (21–56 μeq 1^-1) were observed in lakes above the treeline. Mean relative composition of cations does not differ between July and October; small changes are in the mean relative composition of anions. Acidification of lakes, expressed as a decrease in alkalinity, is 100 μeq l^-1, and is equal to the increase in the sum of sulphate and nitrate. The values of total phosphorus and COD are the lowest in the range of pH 5–6.5. Alkalinity, sulphate, nitrate and pH do not show any trend with time over the last ten years.     

Littoral microcrustaceans (Cladocera and Copepoda) as indices of recovery of a limed water system

A 7 year study (1992–1998) of littoral microcrustaceans (Cladocera and Copepoda) in the watercourse of the River Rore, South Norway, illustrates that qualitative data on cladocerans and copepods are well suited to indicate the recovery of lakes following liming. Eight limed, two acid and two circum neutral reference lakes, were sampled twice a year (June/July and September/October). In the limed lakes, species associated with neutral lakes have become more common, whereas apparently acid-tolerant species have become rarer. In Lake Rore and Lake Syndle, the two largest limed lakes which exhibited a gradual increase in pH, the changes in species composition indicated that these lakes were about to recover. Species composition in Lake Røynelandsvatn, which has reacidified after liming, first reflected improved water quality, then reverted to dominance by acid-tolerant species. In the remaining lakes, the species composition reflects a fauna which has recovered compared with the preliming situation. There is strong evidence, however, that temporary fluctuations in pH have a negative influence on the speed of recovery, confirming the importance of keeping pH stable.     

Influence of dolomitic lime on DOC and DON leaching in a forest soil

The influence of liming on leaching and distribution of dissolved organic carbon (DOC) and nitrogen (DON) in mineral soil was investigated in a leaching experiment with soil columns. Soil samples from separate horizons (O, A and B horizons) were collected from control and limed plots in a field liming experiment in a spruce forest in southern Sweden. The field liming (0.88 kg m^-2) had been carried out 8 years before sampling. To minimize the variation among replicates, soil profiles were reconstructed in the laboratory so that the dry weight was the same for each individual soil horizon regardless of treatment. Two soil column types were used with either the O+A horizons or the O+A+B horizons. One Norway spruce seedling (Picea abies (L.) Karst) was planted in each soil column. Average pH in the leachate water was greater in the limed treatment than in the control treatment (5.0 versus 4.0 for O+A columns and 4.3 versus 3.8 for O+A+B columns). After reaching an approximate steady state, the leaching of DOC was 3--4 times greater from the limed O+A and O+A+B columns than from the corresponding control columns but the leaching of DON increased (3.5 times) only in the limed O+A columns. There was a significant correlation between DOC and DON in the leachates from all columns except for the control O+A+B columns, which indicated a decoupling of DOC and DON retention in the B horizon in the control treatment. This might be explained by a selective adsorption of nitrogen poor hydrophobic compounds (C/N ratio: 32--77) while there was a lower retention of nitrogen rich hydrophilic compounds (C/N ratio: 14--20). Proportionally more hydrophobic compounds were leached from the limed soil compared to the unlimed soil. These hydrophobic compounds also became more enriched in nitrogen after liming so in the limed treatment nitrogen might be adsorbed at nearly the same proportion as carbon, which might explain the fact that there was no decoupling of leached DOC and DON from the B horizon after liming.     

Role of magnesium in combination with liming in alleviating acid-soil stress with the aluminium-sensitive sorghum genotype CV323

An experiment to study the effects of Mg nutrition on root and shoot development of the Al-sensitive sorghum (Sorghum bicolor (L.) Moench) genotype CV323 grown in pots of sandy loam under different acid soil stress is reported. This experiment had a factorial design: four rates of liming were combined with four rates of Mg fertilization. When no Mg was added, the pH of the soil solutions (collected in ceramic cups) increased from 4.0 (unlimed) to 4.2, 4.7 and 5.9 at the increasing rates of liming. After 30 days of growth dry matter yields of the limed treatments were 40%, 115% and 199% higher than that of the unlimed treatment. Without liming and at the highest liming rate, adding Mg did not affect plant biomass significantly. At the two intermediate levels of liming, however, 11.3 mg extra Mg per kg soil increased dry matter yield to the same levels as found at the highest liming rate. Concentrations of Mg in the soil solution rose after Mg was added and fell when lime was added, but adding both Mg and lime increased Mg concentrations in the plant shoots. In plants of the limed treatments, dry matter yield was correlated closely with the Mg concentration in the shoot. This was not so in the unlimed treatment. Furthermore, in the unlimed treatments root development was inhibited, but reduced Mg uptake by the plants resulted mainly from the direct effect of Al- (or H-) ions in the soil solution rather than from impaired root development. It is concluded that Mg fertilization counteracted the interfering effects of Al- and H ions on Mg uptake.     

The diversity of Phaseolus-nodulating rhizobial populations is altered by liming of acid soils planted with Phaseolus vulgaris L. in Brazil

PCR-mediated restriction fragment length polymorphism (RFLP) analysis of the 16S-23S rRNA internally transcribed spacer (ITS) region and the 16S rRNA gene indicated that the rhizobial populations isolated from common bean (Phaseolus vulgaris L.) nodules in the unlimed soil from a series of five lime rates applied 6 years previously to plots of an acidic oxisol had less diversity than those from plots with higher rates of liming. Isolates affiliated with Rhizobium tropici IIB and Rhizobium leguminosarum bv. phaseoli were predominant independent of lime application. An index of richness based on the number of ITS groups increased from 2.2 to 5.7 along the soil liming gradient, and the richness index based on "species" types determined by RFLP analysis of the 16S rRNA gene varied from 0.5 to 1.4. The Shannon index of diversity, based on the number of ITS groups, increased from 1.8 in unlimed soil to 2.8 in limed soil, and, based on RFLP analysis of the 16S rRNA gene, ranged from 0.9 to 1.4. In the limed soil, the subpopulation of R. tropici IIB pattern types contained the largest number of ITS groups. In contrast, there were more R. leguminosarum bv. phaseoli types in the unlimed soil with the lowest pH than in soils with the highest pH. The number of ITS ("strain") groups within R. leguminosarum bv. phaseoli did not change with increased abundance of rhizobia in the soil, while with R. tropici IIB, the number of strain groups increased significantly. Some cultural and biochemical characteristics of Phaseolus-nodulating isolates were significantly related to changes in soil properties caused by liming, largely due to changes in the predominance of the rhizobial species groups.     

Carbon flow in an upland grassland: effect of liming on the flux of recently photosynthesized carbon to rhizosphere soil

The effect of liming on the flow of recently photosynthesized carbon to rhizosphere soil was studied using ¹³CO₂ pulse labelling, in an upland grassland ecosystem in Scotland. The use of ¹³C enabled detection, in the field, of the effect of a 4‐year liming period of selected soil plots on C allocation from plant biomass to soil, in comparison with unlimed plots. Photosynthetic rates and carbon turnover were higher in plants grown in limed soils than in those from unlimed plots. Higher δ¹³C‰ values were detected in shoots from limed plants than in those from unlimed plants in samples clipped within 15 days of the end of pulse labelling. Analysis of the aboveground plant production corresponding to the 4‐year period of liming indicated that the standing biomass was higher in plots that received lime. Lower δ¹³C‰ values in limed roots compared with unlimed roots were found, whereas no significant difference was detected between soil samples. Extrapolation of our results indicated that more C has been lost through the soil than has been gained via photosynthetic assimilation because of pasture liming in Scotland during the period 1990–1998. However, the uncertainty associated with such extrapolation based on this single study is high and these estimates are provided only to set our findings in the broader context of national soil carbon emissions.     

Regulation of bacterioplankton production and biomass in an oligotrophic cleanvater lake--the importance of the phytoplankton community

Estimations of bacterioplankton production and biomass werecarried out in enclosure experiments during two consecutiveyears (1989 and 1990) in oligotrophic clearwater Lake Njupfatet.The lake was limed in November 1989, and the experiments werecarried out both in 1989 (unlimed) and in 1990 (limed). Bags(3001) were manipulated with inorganic phosphorus and nitrogen,organic carbon, and metazoan zooplankton abundance. Both years,bacterial production was stimulated by inorganic nutrients aloneand in combination with organic carbon. However, the increasein bacterial production when inorganic nutrients were addedalone was much stronger in 1990 than in 1989. In 1989. bacterialproduction increased strongly only when inorganic nutrientsand organic carbon were added together. The phytoplankton communitywas dominated by the cyanobacterium Merismopedia tenuis-simaduring 1989, and the phytoplankton biomass increased only slightlywhen receiving inorganic nutrients. In 1990, when the lake hadbeen limed. M.tenuissima had completely disappeared and thephytoplankton community, dominated by Chrysophyceae and Chlorophyceae,responded strongly to additions of inorganic nutrients. Theincreased phytoplankton productivity in 1990 may have resultedin increased release of organic carbon, and this in turn thatthe carbon limitation of bacterioplankton production decreasedfrom 1989 to 1990. Zooplankton had a positive effect on bacterioplanktonproduction in 1989, but no effect in 1990. The loss of bacterialbiomass approximated 60% of the bacterial production in 1989,while in 1990 it almost equalled the bacterioplankton production.     

Effects of liming on rhizosphere chemistry and growth of fine roots and of shoots of sessile oak (Quercus petraea)

Soil acidification can be detrimental to root growth and nutrient uptake, and liming may alleviate such acidification. In the following study, seedlings of sessile oak (Quercus petraea Liebl. M.) were grown in rhizotrons and subjected to liming (L) or gypsum (G) treatments and compared with the control (C). In order to study and interpret the impact of these calcium rich treatments on fine root development and tree growth, the following parameters were assessed: fine root biomass, fine root length, seedling development (height, diameter, leaves), seedling biomass, nutrient content of roots and seedlings, bulk soil and soil solution chemistry and rhizosphere soil chemistry. The results show that liming increased bulk soil pH, exchangeable Mg, Ca and the Ca/Al molar ratio, and decreased exchangeable Al, mainly in the A-horizon. Gypsum had a similar but smaller impact on exchangeable Al, Ca, H⁺ and the Ca/Al molar ratio in the A-horizon, but reacted with depth, so that exchangeable Mn, Mg and Ca were increased in the B-horizon. In the rhizosphere, the general pattern was determined by the treatment effects of the bulk soil. Most elements were more concentrated in the rhizosphere than in bulk soil, except for Ca which was less concentrated after liming or gypsum application. In the B-horizon rhizosphere pH was increased by the treatments (L > G,C) close to the root tips. Furthermore, the length of the zone with a positive root-induced pH increase was greater for the limed roots as compared with both the other treatments. Fine root growth was stimulated by liming (L > G,C) both in terms of biomass and length, whereas specific root length was not obviously affected apart from the indication of some stimulation after liming at the beginning. The live:dead ratio of fine roots was significantly higher in the limed rhizotrons as compared to the control (G not assessed), indicating lower mortality (higher longevity). Shoot growth showed greater lime-induced stimulation (L > G,C) as compared to root growth. As a result the shoot:root ratio was higher in the limed rhizotrons than in the control (L > G,C). Liming induced a higher allocation of P, S, Mg, Ca and K to the leaves, stem and twigs. Gypsum showed similar effects, but was only significant for S. Liming increased the foliar Ca/Al ratio by both increasing foliar Ca and decreasing foliar Al, whereas gypsum did not clearly improve foliar nutrition. This study suggests that a moderate application of lime can be successful in stimulating seedling growth, but that gypsum had no effect on seedling growth. It can be concluded that this lime-induced growth stimulation is directly related to the improved soil fertility status, and the alleviation of Al toxicity and acid stress, resulting in better foliar nutrition. The impact of liming on fine roots, as a consequence, was not limited to a stimulation of the total amount of fine roots, but also improved the root uptake performance. This revised version was published online in June 2006 with corrections to the Cover Date.