Interacting effects of wildfire severity and liming on nutrient cycling in a southern Appalachian wilderness area

Aims

Wilderness and other natural areas are threatened by large-scale disturbances (e.g., wildfire), air pollution, climate change, exotic diseases or pests, and a combination of these stress factors (i.e., stress complexes). Linville Gorge Wilderness (LGW) is one example of a high elevation wilderness in the southern Appalachian region that has been subject to stress complexes including chronic acidic deposition and several wildfires, varying in intensity and extent. Soils in LGW are inherently acidic with low base cation concentrations and decades of acidic deposition have contributed to low pH, based saturation, and Ca:Al ratio. We hypothesized that wildfires that occurred in LGW followed by liming burned areas would accelerate the restoration of acidic, nutrient depleted soils. Because soils at LGW had extremely low concentrations of exchangeable Ca²⁺ and Mg²⁺ dolomitic lime was applied to further boost these cations. We evaluated the effectiveness of dolomitic lime application in restoring exchangeable Ca²⁺ and Mg²⁺ and subsequently increasing pH and Ca:Al ratio of soils and making Ca and Mg available to recovering vegetation.

Methods

Five treatment areas were established: severely burned twice (2000 & 2007) with dolomitic lime application (2xSBL); moderately burned twice with lime application (2xMBL); severely burned twice, unlimed (2xSB); moderately burned once (2000), unlimed (1xMB); and a reference area (REF; unburned, unlimed). In 2008 and 2009, we measured overstory, understory, and ground-layer vegetation; forest floor mass and nutrients; and soil and soil solution chemistry within each treatment area.

Results

All wildfire burned sites experienced substantial overstory mortality. However, understory biomass doubled between sample years on the most recently burned sites due to the rapid regrowth of ericaceous shrubs and prolific sprouting of deciduous trees. Burning followed by lime application (2xSBL and 2xMBL) significantly increased shallow soil solution NO₃-N, but we found no soil solution NO₃-N response to burning alone (2xSB and 1xMB). Surface soil base saturation and exchangeable Ca²⁺ were significantly affected by liming; Ca²⁺ concentrations were greater on 2xMBL and 2xSBL than 2xSB, 1xMB and REF. There was a smaller difference due to moderate burning along with greater soil Ca²⁺ on 1xMB compared to REF, but no difference between 2xSB and REF. Surface and subsurface soil exchangeable Al³⁺ were lower on 2xSBL than 2xSB, 2xMBL, 1xMB, and REF. Liming decreased soil acidity somewhat as surface soil pH was higher on the two burned sites with lime (pH?=?3.8) compared to 2xSB without lime (pH?=?3.6).

Conclusions

Liming resulted in decreased soil Al³⁺ on 2xSBL coupled with increased soil Ca²⁺ on both 2xSBL and 2xMBL, which improved soil Ca/Al ratios. However, the soil Ca/Al ratio response was transitory, as exchangeable Al³⁺ increased and Ca/Al ratio decreased over time. Higher lime application rates may be necessary to obtain a substantial and longer-term improvement of cation-depleted soils at LGW.     

Influence of pH,exchangeable aluminium and 0.02M CaCl2-extractable aluminium on the growth and nitrogen-fixing activity of white clover (Trifolium repens) in some New Zealand soils

The effects of soil acidity on the growth and N₂-fixing activity of white clover in seven acid topsoils and subsoils of New Zealand were investigated using a glasshouse experiment.The application of phosphate (Ca(H₂PO₄)₂) to the soils resulted in very large increases in white clover growth on all soils. The application of phosphate, as well as increasing P supply, also decreased 0.02M CaCl₂-extractable Al levels, but had little effect on exchangeable Al levels.Where adequate phosphate was applied, increasing rates of lime (CaCO₃) resulted in increased plant growth on most soils. N₂[C₂H₂]-fixing activity was increased by the first level of lime for one soil, but generally remained approximately constant or declined slightly at higher rates of lime. Up to the point of maximum yield, white clover top weight was more highly correlated with 0.02M CaCl₂-extractable soil Al than with exchangeable Al or pH. At pH values greater than 5.5, plant yield declined on some soils, apparently because of Zn deficiency. The data suggest that white clover is unlikely to be affected by Al toxicity at 0.02M CaCl₂-extractable Al levels of less than about 3.3 g g^–1. However, there were differences between soils in apparent plant tolerance to 0.02M CaCl₂-extractable Al, which appeared to be caused by differing C levels in the 0.02M CaCl₂ extracts.     

The effect of level of CaCO3, inoculation and lime pelleting on the nodulation and growth of beans in five acid soils

Summary The effect of level of CaCO₃, inoculation and lime pelleting on the nodulation, dry matter yield and % N content of common bean plants (Phaseolus vulgaris) grown in five acid soils was investigated in a greenhouse study. The soils represented a range in pH from 3.9 to 5.1, in exchangeable Al from 0.0 to 4 meq/100 gm, in exchangeable Mn from 0.35 to 2.32 me/100 gm, and in %C from 0.69 to 5.60.Nodule weight decreased with increasing %C and for the soil with highest %C (5.60) no nodules were observed. In soils with low organic matter and low exchangeable Al and Mn, inoculation increased nodule weight, dry matter yield and %N especially at the lowest pH level. Where the seeds were not inoculated, nodule weight and dry matter yield increased with soil pH. No such increases were observed where the seeds were inoculated. There was no apparent advantage in lime pelleting in such soils.In soils with low organic matter content and with substantial amounts of Al and/or Mn, liming increased nodule weight and dry matter yield, and decreased exchangeable Al and/or Mn. Lime pelleting was superior to mere inoculation in increasing nodule weight particularly at low lime rates.In soils with relatively high organic matter content, nodulation was very low or none at all. Low lime rates had little effect on exchangeable Al and Ca and dry matter yield. Higher lime rates, however, decreased exchangeable Al and dry matter yield but increased exchangeable Ca.     

Differential acidity tolerance of tropical legumes grown for green manure in acid sulfate soils

The growth of four tropical legumes (Cajanus cajan, Sesbania aculeata, S. rostrata, and S. speciosa) used as green manures in the tropics was studied in a glasshouse experiment. Two acid sulfate soils (Typic Sulfaquept, Bang Pakong Series; and Sulfic Tropaquept, Rangsit Series) were adjusted to four pH levels: 3.8 or 4.0 (original soil pH), 4.5, 5.5, and 6.5 (amended with lime). Dry weight was determined 49 days after sowing. Concentrations of N, P, K, Ca, Mg, Fe, Mn, and Al were also determined in aerial plant parts at harvest.The legumes responded differently to soil acidity and liming, but not to soil type. Cajanus cajan had the highest biomass production, followed by S. aculeata, S. rostrata and S. speciosa, in this order. The N concentration closely paralleled biomass production, suggesting that the growth of symbiotic rhizobia and nodulation were perhaps more susceptible to soil acidity than were the host plants. Liming to pH 5.5–6.0 was recommended for the legumes' growth based on the quadratic relationships between dry-matter yield and soil pH. In the unlimed soils, the Ca concentration in C. cajan and S. aculeata (0.32%) was twice as high as that in the two low-yielding legumes (0.15%). Furthermore, plant Ca increased exponentially (or quadratically in case of S. speciosa) as lime additions increased. It was estimated that for adequate growth, the Ca requirement in the shoot dry matter was approximately: C. cajan 1.2% Ca, S. aculeata 0.8%, S. rostrata 0.6%, and S. speciosa 0.4%. In contrast with Ca, the concentration of Fe, and to a lesser extent Mn, was significantly lower in C. cajan and S. aculeata than in S. rostrata and S. speciosa. The ratio of Ca to Al in plant tops was used to characterize plant tolerance to soil acidity, and to quantify the critical Al concentration in the plants. It appears that 90% maximum growth was attained only when Ca/Al was 150 for C. cajan and S. speciosa, 200 for S. rostrata, and 300 for S. aculeata. Cajanus cajan tolerated up to 80 mg Al kg^-1 in the shoot dry matter, whereas significant growth reduction occurred in the Sesbania species at levels > 30 mg Al kg^-1.     

Effect of liming on spore germination,germ tube growth and root colonization by vesicular-arbuscular mycorrhizal fungi

Summary The effect of soil acidity on spore germination, germ tube growth and root colonization of vesicular-arbuscular mycorrhizal (VAM) fungi was examined using a Florida Ultisol. Soil samples were treated with 0, 4, 8 and 12 meq Ca/MgCO₃/100 g soil and each lime level received 0, 240, and 720 ppm P as superphosphate. Corn (Zea mays L.) was planted in the soil treatments, inoculated with eitherGlomus mosseae orGigaspora margarita spores and grown for 31 days. Acid soil inhibits mycorrhizal formation byG. mosseae through its strong fungistatic effect against the spores. The dolomitic lime increased mycorrhizal formation by both fungal species.G. margarita is much less sensitive to acidic conditions thanG. mosseae. Al ions are a very important component of the fungistatic property against the VAM symbiosis. VAM fungus adaptation may be important for plants growing on infertile acid soils if soil inoculation with these fungi is to contribute significantly to low-input technology for tropical agricultural systems.     

Effect of lime and phosphorus applications on concentrations of available nutrients and on P,Al and Mn uptake by two pasture legumes in an acid soil

Summary Effects of increasing rates of lime and phosphorus addition on concentrations of available nutrients in soil and on P, Al and Mn uptake by two pasture legumes, lotus (Lotus pedunculatus Cav.) and white clover (Trifolium repens L.), were studied in a pot experiment using a highly leached acid (pH 4.2) soil.Liming resulted in an increase in exchangeable Ca and thus in percentage base saturation, with concomitant decreases in levels of exchangeable Al, Fe and Mn. The relationship between exchangeable Ca and Al was linear and negative with a gradient of almost unity. Liming had no consistent effect on measured CEC values. Increasing lime rates significantly reduced concentrations of Mg, K and Na in saturation paste extracts but had no effect on exchangeable Mg, K and Na levels.With increasing lime additions, available phosphate indicesviz water soluble, resin-, Morgan-and Williams-extractable all decreased significantly, Truog-extractable was unaffected, while Brayextractable generally increased. Fractionation revealed that lime additions caused a decrease in easily soluble, Fe-bound and to a lesser extent Ca-bound phosphate fractions, had no effect on reductant soluble phosphate, but resulted in an increase in the Al-bound fraction. P uptake and yield of both legumes increased with lime and P additions.Correlations between available phosphate indices and yield of both legumes were weak or nonsignificant. However, high, significant positive correlation coefficients were found between available phosphate and plant uptake of P. Indices of available Al and Mn were not generally significantly correlated with plant uptake of Al or Mn but significant negative correlations were found between available Al and Mn and yield of both species.     

Effects of phosphate fertilization,lime amendments and inoculation with VA-mycorrhizal fungi on soybeans in an acid soil

Soybeans [Glycine max (L.) Merr. cv. Essex] were grown in nonsterile acid (pH. 5.2) infertile Wynnville silt loam (Glossic Fragiudult) in a glasshouse. The effects of P fertilization and lime were determined by inoculation with two VAM-fungi (VAMF): Glomus fasciculatum (Gf) and Glomus etunicatum (Ge). An important factor affected by the interaction between applied lime (soil acidity), applied P, and VAMF inoculation was the soil Al. Five application rates of P as KH₂PO₄ and three rates of lime were tested. Potassium was equalized with KCl (muriate of potash). P-efficiency (g seed/mg P kg^-1 soil) by vesicular-arbuscular mycorrhiza (VAM) was maximal at 20 mg P kg^-1 soil at all lime and VAMF treatments. VAMF inoculation increased plant survival and protected the soybeans from leaf scorch, thereby substituting for the effects of lime and P. The Ge inoculum was superior in ameliorating leaf scorch in the nonlimed soil. The Gf inoculum required more lime and P than the Ge inoculum to increase seed yield relative to the noninoculated controls containing only native VAMF. Both inocula increased root Al uptake and extractable soil Al in the acid soil without apparent adverse effects on root or shoot. The ability of the VAMF inocula to enhance the efficiency of applied P and decrease seed Cl concentration was increased by lime. Seed yield (Y) was negatively related to seed Cl concentration (X) where Y=aX^-b. Both VAMF inoculation and lime application reduced this negative relationship and may have increased the tolerance to both Cl and soil Al.     

The effect of phosphate rock dissolution on soil chemical properties and wheat seedling root elongation

Soils of the Appalachian region of the United States are acidic and deficient in P. North Carolina phosphate rock (PR), a highly substituted fluoroapatite, should be quite reactive in these soils, allowing it to serve both as a source of P and a potential ameliorant of soil acidity. An experiment was conducted to evaluate the influence of PR dissolution on soil chemical properties and wheat (Triticum aestivum cv. Hart) seedling root elongation. Ten treatments including nine rates of PR (0, 12.5, 25, 50, 100, 200, 400, 800, and 1600 mg P kg^-1) and a CaCO₃ (1000 mg kg^-1) control were mixed with two acidic soils, moistened to a level corresponding to 33 kPa moisture tension and incubated for 30 days. Pregerminated wheat seedlings were grown for three days in the PR treated soils and the CaCO₃ control. Root length was significantly (P<0.05) increased both by PR treatments and CaCO₃, indicating that PR dissolution was ameliorating soil acidity. The PR treatments increased soil pH, exchangeable Ca, and soil solution Ca while lowering exchangeable Al and 0.01 M CaCl₂ extractable soil Al. Root growth in PR treatments was best described by an exponential equation (P<0.01) containing 0.01 M CaCl₂ extractable Al. The PR dissolution did not reduce total soil solution Al, but did release Al complexing anions into soil solution, which along with increased pH, shifted Al speciation from toxic to nontoxic forms. These results suggest that North Carolina PR should contribute to amelioration of soil acidity in acidic, low CEC soils of the Appalachian region.     

Relationships among foliar chemistry, foliar polyamines, and soil chemistry in red spruce trees growing across the northeastern United States

Forest trees are constantly exposed to various types of natural and anthropogenic stressors. A major long-term goal of our research is to develop a set of early physiological and biochemical markers of stress in trees before the appearance of visual symptoms. Six red spruce (t Picea rubens Sarg.) stands from the northeastern United States were selected for collection of soil and foliage samples. All of the chosen sites had soil solution pH values below 4.0 in the Oa horizon but varied in their geochemistry. Some of these sites were apparently under some form of environmental stress as indicated by a large number of dead and dying red spruce trees. Samples of soil and needles (from apparently healthy red spruce trees) were collected from these sites four times during a two-year period. The needles were analyzed for perchloric acid-soluble polyamines and exchangeable inorganic ions. Soil and soil solution samples from the Oa and B horizons were analyzed for their exchange chemistry. The data showed a strong positive correlation between Ca and Mg concentrations in the needles and in the Oa horizon of the soil. However, needles from trees growing on relatively Ca-rich soils with a low exchangeable Al concentration and a low Al:Ca soil solution ratio had significantly lower concentrations of putrescine and spermidine than those growing on Ca-poor soils with a high exchangeable Al concentration and a high Al:Ca soil solution in the Oa horizon. The magnitude of this change was several fold higher for putrescine concentrations than for spermidine concentrations. Neither putrescine nor spermidine were correlated with soil solution Ca, Mg, and Al concentrations in the B horizon. The putrescine concentrations of the needles always correlated significantly with exchangeable Al (r²=0.73, t p0.05) and soil solution Al:Ca ratios (r²=0.91, t p0.01) of the Oa horizon. This suggests that in conjunction with soil chemistry, putrescine and/or spermidine may be used as a potential early indicator of Al stress before the appearance of visual symptoms in red spruce trees.     

Influence of organic matter on the availability of certain elements to barley seedlings grown by a modified neubauer method

Summary The influence of organic matter on the availability of 17 elements (Na, K, Cs¹³⁷, Mg, Ca, Sr, Ba, N, P, B, Cu, Zn, Fe, Mn, Mo, Al, and Si) to barley seedlings grown by a modified Neubauer technique was determined. Three different soils that were treated with dry ground mustard spinach leaves (1 g/100 g soil) and incubated for various lengths of time (0, 1, 2, 5, 9, 13, and 17 weeks) in moist condition before cropping were used for this study.The addition of organic matter to the soils increased the plant yields. The average N and K concentrations were consistently increased in the plants grown in soils with added organic matter. The average concentration of B, P, Na, Mg, Sr, Ba, and Si were almost consistently decreased in the plants. The average contents of Cu, Zn, Fe, Mn, Mo, Ca, and Al varied with the soil types and precropping incubation time. The average Cs¹³⁷ contents of the plants were reduced considerably by the addition of organic matter to the soils. The reduction of Cs¹³⁷ contents ranged from 29 to 75 per cent, depending on the pre-cropping incubation time and soil type. The main factors causing this reduction were considered to be microbial immobilization, ion antagonism by K, carbohydrate dilution, and the state of decomposition and the kind of organic matter added to the soils.     

Monitoring uptake and contents of Mg,Ca and K in Norway spruce as influenced by pH and Al,using microprobe analysis and stable isotope labelling

In a model system using intact spruce trees (Picea abies [L.] Karst.) we followed the path of magnesium, calcium and potassium during uptake into the root and during long-range transport into the shoot, by multiple stable isotope labelling. The roots of two- and three-year-old spruce trees originating from soil culture were removed from the soil and, in part or in toto, exposed to labelling solutions containing the stable isotopes ²⁵Mg or ²⁶Mg, ⁴¹K and ⁴²Ca or ⁴⁴Ca. Optical-emission-spectroscopy (ICP-OES) of plant fractions and labelling solutions was combined with the quantitative analysis of stable isotope ratios in sections of shock frozen, cryosubstituted material using the laser-microprobe-mass-analyser (LAMMA). This combination allowed us to distinguish, both in bulk samples and on the cellular level between (i) the fraction of elements originally present in the plant before the start of the labelling, (ii) the material taken up from the labelling solution into the plant and (iii) any material released by the plant into the labelling solution.In single-root labelling experiments, roots of three-year-old spruce trees, grown in nursery soil, were exposed to various pH conditions. The exchange of Mg and Ca with the labelling solution was nearly 100% in the cell walls of the mycorrhized finest roots. This exchange was only slightly affected by a step down to pH 3.5. The absolute Mg and Ca content in the cell walls was moderately reduced by incubation at pH 3.5 and strongly reduced in the presence of Al at this pH. After a pH 3.5 and 2 mM Al treatment we found Al in the xylem cell walls and the cortex cell lumina at elevated concentrations. To analyse the combined effect of high Al and high proton concentrations on the long-range transport, we used a split-root system. The root mass of an intact two-year-old spruce tree, grown in mineral soil, was divided into even parts and both halves incubated in solutions with two sets of different stable isotopes of Mg and Ca (side A: no Al, ²⁵Mg and ⁴²Ca; side B: +Al, ²⁶Mg and ⁴⁴Ca) and ⁴¹K on both sides. We observed a large uptake of Mg, Ca and K into the plant and a pronounced release. The net uptake of all three elements was lower from the Al-doted solution. In cross-sections of the apical shoot we found after seven-day labelling period about 60–70% of the Mg and Ca and 30% of the K content in the xylem cell walls originating from both labelling solutions. The clear majority of the Mg and Ca label originated from the Al-doted side.     

Effects of liming on soil chemical characteristics and grass growth in laboratory and long-term field-amended soils

Summary The influence of liming on soil solution composition was compared in two laboratory amended soils and one field amended soil. In the laboratory study, soil solutions were sampled by miscible displacement at intervals of 1 and 10 weeks after liming. In addition to increases in pH and Ca, there were large reductions in the concentrations of Mg, K, Na, Si and Mn. Solution concentration of free Al decreased with liming; however, organically complexed Al increased, as did soluble organic matter. Liming also stimulated mineralization of N as indicated by increased solution NO₃ levels. The field amended soils were obtained from a long-term cutting trial investigating the effects of lime on pasture. Despite the passage of a 16-year interval since application, the effects of lime on soil solution characteristics were still clearly evident and generally consistent with those observed in the laboratory study. Estimated leaching losses of Ca from limed soil were relatively low, amounting to 12%, 27% and 44% of the 4.2, 8.4 and 12.5 t lime ha⁻¹ applied, respectively. The results suggest that, in Eastern Ireland, a lime treatment would maintain and elevated pH and would influence the avialability and mobility of plant nutrients for some decades following application.     

Dry matter yield,root traits,and nodule occupancy of lucerne and Caucasian clover when grown in acidic soil with high aluminium concentrations

Aims and methods

Lucerne and Caucasian clover dry matter were measured in response to recommended lime and capital P inputs for six years in an acidic soil in the New Zealand high country. The initial three years of the field experiment indicated successful establishment and persistence of both legumes. Lucerne dry matter (DM) yield was up to 4 t/ha/yr in this period and higher than Caucasian clover yields. However, a lack of persistence of lucerne was apparent from this point forward compared with Caucasian clover which produced 7.7 t DM/ha in Year 6. An experiment using tubes of soil was used to investigate whether differences in root traits, nodulation and nodule occupancy were responsible for the differences observed in field persistence over time.

Results

These showed that when rhizobia inoculant was added, the fine root length of Caucasian clover was unaffected (R² = 0.14) by aluminium (Al) content of the soil. In contrast, fine root growth of lucerne was suppressed (R² = 0.79) by the soil Al content. Nodulation of Caucasian clover was unaffected by soil pH or Al when the rhizobia inoculant was provided which suggests the viability of the commercial genotype ICC148 in this soil with a pH of 5.5 and Al ca. 7 mg/kg soil. For lucerne, the maximum nodulation score of 7.3 occurred with 2 t/ha of lime added (soil pH ca.6, Al ca. 0.3 mg/kg) plus inoculant.

Conclusions

This suggests an Al toxic threshold of <1.0 mg Al/kg soil for effective lucerne nodulation. From the lucerne nodules, eight naturalized strains of Ensifer meliloti were identified. In contrast, only one R. leguminosarum strain was detected in the Caucasian clover nodules. The competition between those rhizobia genotypes may negatively affect the efficiency of biological nitrogen fixation in lucerne. Therefore, the lack of genetic diversity of R. leguminosarum bv. trifolii in New Zealand soils might be an advantage especially if the commercial strain is acid soil tolerant.

     

Polyphenols as Regulators of Plant-litter-soil Interactions in Northern California's Pygmy Forest: A Positive Feedback?

The convergent evolution of polyphenol-rich plant communities has occurred on highly acidic and infertile soils throughout the world. The pygmy forest in coastal northern California is an example of an ecosystem on an extremely infertile soil that has exceptionally high concentrations of polyphenols. Many negative feedbacks have been identified whereby plants degrade fertile soils through production of polyphenol-rich litter, sequestering soil nutrients into unavailable form and creating unfavorable conditions for seed germination, root growth, and nutrient uptake. But in the context of plant-litter-soil interactions in ecosystems adapted to soils that are inherently acidic and infertile (such as the pygmy forest), there are also many positive feedbacks that result from polyphenol production. By inhibiting decomposition, polyphenols regulate the formation of a mor-humus litter layer, conserving nutrients and creating a more favorable medium for root growth. Polyphenols shift the dominant pathway of nitrogen cycling from mineral to organic forms to minimize potential N losses from the ecosystem and maximize litter-N recovery by mycorrhizal symbionts. Polyphenol complexation of Al, Mn and Fe reduce potential Al toxicity and P fixation in soil. Polyphenols regulate organic matter dynamics, leading to the accumulation of organic matter with cation exchange capacity to minimize leaching of nutrient cations. Humic substances derived from polyphenolic precursors coat rhizosphere soil surfaces, improving physical and chemical conditions for root growth and nutrient cycling. Although their long-accepted adaptive value for antiherbivore defense is now in doubt, polyphenol alteration of soil conditions and regulation of nutrient cycling illustrate how fitness can be influenced by the extended phenotype in plant-litter-soil interactions.     

The role of ammonia volatilization in controlling the natural 15N abundance of a grazed grassland

Although the variation in natural ¹⁵N abundance in plants and soils is well characterized, mechanisms controlling N isotopic composition of organic matter are still poorly understood. The primary goal of this study was to examine the role of NH₃ volatilization from ungulate urine patches in determining ¹⁵N abundance in grassland plants and soil in Yellowstone National Park. We additionally used isotopic measurements to explore the pathways that plants in urine patches take up N. Plant, soil, and volatilized NH₃¹⁵N were measured on grassland plots for 10 days following the addition of simulated urine. Simulated urine increased ¹⁵N of roots and soil and reduced ¹⁵N of shoots. Soil enrichment was due to the volatilization of isotopically light NH₃. Acid-trapped NH₃¹⁵N increased from –28 (day 1) to –0.3 (day 10), and was lighter than the original urea-N added (1.2). A mass balance analysis of urea-derived N assimilated by plants indicated that most of the N taken up by plants was in the form of ammonium through roots. However, isotope data also showed that shoots directly absorbed ¹⁵N – depleted NH₃-N that was volatilized from simulated urine patches. These results indicate that NH₃ volatilization from urine patches enriches grassland soil with ¹⁵N and shoots are a sink for volatilized NH₃, which likely leads to accelerated cycling of excreted N back to herbivores.     

The influence of soil chemistry on fine root aluminum concentrations and root dynamics in a subalpine Spodosol,Washington State,USA

The relationship between root Al concentrations and Al fractions in the soil solution was examined in a mature Abies amabilis ecosystem in the Cascade Range of Washington State. The naturally acidic soils in these ecosystems lead to high concentrations of aqueous Al in soil solutions and contribute to the biocycling of Al by the A. amabilis/T. mertensiana stand. Root concentrations of Al were very closely related to aqueous Al³⁺ activities, but poorly correlated with total aqueous Al concentrations. The solution Al/Ca molar ratios followed a seasonal cycle with low values during the fall and high values during the spring. Ratios remained <1 throughout the year in the Oa horizon while they varied between 2 and 14 in the E and Bhs horizons. The vertical distribution of roots and the mortality of fine roots may be linked to the soil solution Al/Ca ratio. Root cation exchange capacity ranged between 180 and 225 mol g^-1 and the exchangeable Al fraction represented from 12–17% of the total Al content in the root. Evidence for solid-phase co-precipitates of Al with PO₄ and oxalate was indicated from selective dissolution of the root tissue. Sufficient quantities of PO₄ and oxalate exist in the roots to tie up 20–40% of the Al present in the roots of the Oa and E horizons, but only 9% of that present in the Bhs horizon. Species differences in the distribution of Al between the above-ground and below-ground components may be dictated by these retention processes in the fine roots.     

Vesicular-arbuscular mycorrhizal effectiveness in an acid soil amended with fresh organic matter

A greenhouse investigation was undertaken to determine the influence of fresh organic matter on the formation and functioning of vesicular-arbuscular mycorrhizal symbiosis in Leucaena leucocephala grown in an acid aluminum-rich ultisol. In soil not amended with fresh organic matter or lime, plants failed to grow. Mycorrhizal infection level, mycorrhizal effectiveness measured in terms of pinnule P content of L. leucocephala leaves and dry matter yield of the legume increased with increase in fresh organic matter. Although VAM colonization level and dry matter yield of L. leucocephala were significantly higher if the test soil was limed (7.2 cmole OH^–) than if amended with fresh organic matter, the latter was as effective as lime in off-setting the detrimental effect of aluminum on mycorrhizal effectiveness. The lower mycorrhizal colonization level and the lower dry matter yield noted in the soil treated with fresh organic matter appears to be related to the inadequacy of Ca in the soil amended with fresh organic matter. These observations are supported by the low calcium status of soil and plant tissues in the absence of lime. It is concluded that while fresh organic matter, in appropriate amounts, could protect sensitive plants and VAM symbiosis against Al toxicity in acid soils, maximum mycorrhizal inoculation effects are not likely to be attained unless the soils are also amended with Ca.Contribution from Hawaii Institute of Tropical Agriculture and Human Resources Journal Series No 3740.     

Aluminium Uptake and Translocation in Al Hyperaccumulator Rumex obtusifolius Is Affected by Low-Molecular-Weight Organic Acids Content and Soil pH

Background and Aims

High Al resistance of Rumex obtusifolius together with its ability to accumulate Al has never been studied in weakly acidic conditions (pH > 5.8) and is not sufficiently described in real soil conditions. The potential elucidation of the role of organic acids in plant can explain the Al tolerance mechanism.

Methods

We established a pot experiment with R. obtusifolius planted in slightly acidic and alkaline soils. For the manipulation of Al availability, both soils were untreated and treated by lime and superphosphate. We determined mobile Al concentrations in soils and concentrations of Al and organic acids in organs.

Results

Al availability correlated positively to the extraction of organic acids (citric acid < oxalic acid) in soils. Monovalent Al cations were the most abundant mobile Al forms with positive charge in soils. Liming and superphosphate application were ambiguous measures for changing Al mobility in soils. Elevated transport of total Al from belowground organs into leaves was recorded in both lime-treated soils and in superphosphate-treated alkaline soil as a result of sufficient amount of Ca available from soil solution as well as from superphosphate that can probably modify distribution of total Al in R. obtusifolius as a representative of “oxalate plants.” The highest concentrations of Al and organic acids were recorded in the leaves, followed by the stem and belowground organ infusions.

Conclusions

In alkaline soil, R. obtusifolius is an Al-hyperaccumulator with the highest concentrations of oxalate in leaves, of malate in stems, and of citrate in belowground organs. These organic acids form strong complexes with Al that can play a key role in internal Al tolerance but the used methods did not allow us to distinguish the proportion of total Al-organic complexes to the free organic acids.     

Rapid evaluation of sorghum for aluminum tolerance

Two screening methods tested in this study were based on the observation that roots of freshly germinated seeds exhibit their relative tolerance to Al during the first 48 hours of growth. Sorghum (Sorghum bicolor L. Moench) varieties were evaluated using toxic/nontoxic soil pairs in petri dishes and toxic/nontoxic solution pairs in a flowing aqueous system. The soils had 0, 61, and 72% Al saturation and the solutions had 0, 1.85 and 3.70 M l^–1 Al from AlCl₃ and 0.25 mM l^–1 Ca from Ca(NO₃)₂. Relative root lengths in both systems correctly differentiated 13 genotypes of known Al tolerance into tolerant and intolerant groups. Twenty six other sorghums genotypes were also screened using genotypes of known Al reactions as checks. The soil with 72% Al saturation and the 1.85 M l^–1 Al solution gave the highest correlation between the two systems and both effectively arrayed sorghums of known and unknown tolerance. For routine screening the authors prefer the soil system for its simplicity, efficiency, and use of a natural growth medium.Journal paper 11637. Purdue University Agricultural Experiment Station, Lafayette, Indiana. Contribution from the Agronomy Department.     

Contributions of three subsystems of a freshwater marsh to total bacterial secondary productivity

Rates of bacterial production were measured in the water column, on the surface of plant detritus, and in the surface sediments of a freshwater marsh in the Okefenokee Swamp, Georgia, USA. Bacterioplankton production rates were not correlated with several measures of quantity and quality of dissolved organic matter, including an index of the relative importance of vascular plant derivatives. Bacterioplankton productivity was high (mean: 63 g C liter^–1 day^–1) compared with rates reported for other aquatic ecosystems. Somewhat paradoxically, bacterial productivity on plant detritus (mean: 13 g C g^–1 day^–1) and sediments (mean: 15 g C g^–1 day^–1) was low relative to other locations. On an a real basis, total bacterial productivity in this marsh ecosystem averaged 22 mg C m^–2 day^–1, based on sample dates in May 1990 and February 1991. Marsh sediments supported the bulk of the production, accounting for 46% (May) and 88% (February) of the total. The remainder was contributed approximately equally by bacteria in the water column and on accumulated stores of plant detritus. Send offprint requests to: M. A. Moran.