Inability to solubilize phosphate in limestone soils—key factor controlling calcifuge habit of plants

Germination, seedling establishment and growth of calcifuge plants in Swedish limestone soils of Archean and Ordovician age were studied. As previously demonstrated for Viscaria vulgaris, establishment of Rumex acetosella and Silene rupestris did not succeed unless CaHPO₄ (at the rate of 10 mmol dm^-3 of soil) was supplied. Growth of Deschampsia flexuosa was enhanced by phosphate addition, whereas establishment success of Jasione montana was poor, regardless of phosphate treatment. Establishment and growth in an acidic gneiss soil, used as a reference for the species studied, was good. Total, total inorganic, exchangeable, and soil solution P were considered in all soils and treatments. It is proposed that the calcifuge behaviour of plants is quite often caused by inability to solubilize the native phosphate of limestone soils.     

A New Approach to Understanding the Calcifuge Habit of Plants

Growth rate of the calcifuge plants Carex pilulifera, Deschampsiaflexuosa, Holcus mollis, Luzula pilosa, Nardus stricta, andVeronica officinalis, transplanted into an Ordovician limestonesoil of pH 8, increased by two to three times on addition of5 mol m^-3 of CaHPO₄ compared to untreated conditions. For Galiumsaxatile, however, P treatment was lethal and growth was possibleonly in soil supplied with Fe(III) citrate, which had littleor no effect on growth of the other six species. Phosphate treatmentof the limestone soil greatly increased plant uptake of P, whereasP concentration of shoots from untreated soil was very low andprobably highly deficient, compared to plants of field siteorigin. From this and two other recent studies it is now possibleto conclude that the calcifuge habit of plants, at least underclimatic conditions prevailing in northern Europe, is most oftendue to an inability of such plants to render the native phosphateof limestone soils available to plant uptake. Out of ten calcifugespecies tested, only one exception to this rule was identified.Copyright1994, 1999 Academic Press Carex pilulifera, Deschampsia flexuosa, Galium saxatile, Holcus mollis, Luzula pilosa, Nardus stricta, Veronica officinalis, calcifuge plants, phosphorus, iron, limestone soil, limiting factors     

Mineral Nutrient Limitations of Calcifuge Plants in Phosphate Sufficient Limestone Soil

Twelve species of calcifuge plants were grown in an Ordovician-limestone soil with and without phosphate amendment, as well as in an acid silicate soil of their natural habitat. Phosphate treatment of the limestone soil raised the P concentrations of the plant biomasses to levels within sufficiency ranges reported for cultivated plants and productivity usually increased two- to five-fold. Out of twelve species studied,Scleranthus perenniswas unable to survive in the limestone soil unless treated with phosphate, whereas growth and general performance ofGalium saxatilewas impaired by phosphate additions. Biomass dilution effects on micro-nutrients, but usually not on macronutrients, were recorded as a result of the phosphate treatment. Dilution of Mn was most distinct and Fe was least distinct. However, no foliar symptoms clearly assignable to Mn deficiency were observed. Symptoms of foliar chlorosis, reminiscent of Fe deficiency, developed inGalium saxatile, Carex piluliferaandVeronica officinalis. InC.pilulifera, but not inV.officinalis, chlorosis was accompanied by decreasing foliar Fe concentrations.     

Potential Nitrification as an Indicator of Preferential Uptake of Ammonium or Nitrate by Plants in an Oak Woodland Understorey

The preferences of some woodland understorey species for ammoniumand nitrate were investigated by measuring the potential nitrification(conversion of ammonium to nitrate) in the rhizosphere comparedwith the bulk soil. Less acid-tolerant species, which usuallyprefer nitrate or a mixture of ammonium and nitrate in hydroponicculture, should have a higher potential nitrification in therhizosphere compared to the bulk soil due to a low uptake ofammonium (since ammonium is relatively immobile). Acid-tolerantspecies should have a high uptake of ammonium and thereby loweror equal potential nitrification in the rhizosphere comparedto the bulk soil. The hypothesis was tested in a field investigationof five understorey herb species, Deschampsia flexuosa, Convallariamajalis, Poa nemoralis, Geum urbanum andAegopodium podagrariaperformed in oak forests in southern Sweden. Overall, the twoless acid-tolerant species, Geum urbanum and Aegopodium podagraria,had high potential nitrification in the rhizosphere comparedto the bulk soil (indicating a relatively low uptake of ammonium),whilst the acid tolerant species, Deschampsia flexuosa andConvallariamajalis , had approximately equal potential nitrification inthe rhizosphere compared to the bulk soil (indicating a relativelyhigh uptake of ammonium). In the case of Poa nemoralis, a specieswhich grows in both acid and less acid soils, we found the potentialnitrification in the rhizosphere and in the bulk soil to besimilar at low inorganic nitrogen concentrations, but the difference(rhizosphere > bulk) increased when nitrification in thebulk soil was enhanced (i.e. when the nitrogen availabilityincreased). The potential nitrification in the bulk soil variedbetween 0 and 16 nmol g^-1h^-1and was positively correlated withpH. When species occurred at the same site, the potential nitrificationin the bulk soil tended to be lower for the acid tolerant species.Despite a large variation in potential nitrification, the methodoffers a possibility of measuring the preference of plants forammonium/nitrate in a soil system, under natural conditions.Copyright 2000 Annals of Botany Company Ammonium uptake, nitrate uptake, nitrogen preference, potential nitrification, rhizosphere, Deschampsia flexuosa, Convallaria majalis, Poa nemoralis, Geum urbanum, Aegopodium podagraria     

Growth and mineral nutrition of plant species from clearings on different horizons of an iron-humus podzol profile

Summary Four plant species of woodland clearings — Avenella flexuosa, Chamaenerion angustifolium, Rumex acetosella, and Senecio sylvaticus — were grown on five different horizons of an iron-humus podzol profile. Reasonable growth was achieved only on the A₀ horizon, which was rich in organic matter. The growth reduction on the other horizons was correlated with low concentrations of manganese and phosphorus in soils and plant organs. The restriction of so-called acidophilous species to acid soils is interpreted as meaning that they have a need for a high supply of manganese.     

Differences in Iron Nutrition Strategies of Two Calcifuges,Carex piluliferaL. andVeronica officinalisL.

Veronica officinalisandCarex pilulifera, widespread calcifugeplants in Europe, were cultivated in acid and calcareous soilsto study differences in Fe aquisition strategies indicated inprevious studies. The experiments were performed in a computer-controlledglasshouse at a soil solution moisture content of 50–60%water holding capacity; additional light was supplied at 70W m^-2if ambient light was <100 W m^-2between 0600 and 1800h.Both species developed chlorosis when grown in the calcareoussoil.C. piluliferaproved unable to translocate sufficient amountsof Fe to the leaves when cultivated in calcareous soil, butmuch Fe accumulated in, and especially as a precipitate on thesurface of roots. In contrast,V. officinalistended to increaseFe taken up into the leaves of plants grown on calcareous soil,but a much greater proportion of the leaf tissue Fe was accumulatedas less active forms not extracted by Fe complexing agents,e.g. 1,10-phenanthroline, than was the case in acid-soil grownplants. Considerably less Fe was accumulated in the root biomassofV. officinaliscompared toC. pilulifera.It is concluded thatchlorosis inC. piluliferais related to insufficient Fe uptakein the leaves, whereas leaf immobilization of Fe in physiologicallyless active forms is the problem inV. officinalis. Iron; chlorosis; calcifuge; iron immobilization; leaf tissue; fractionation; Carex pilulifera; Veronica officinalis     

Arsenate tolerance in Agrostis castellana and Agrostis delicatula

Experiments were made to elucidate why many field-layer plants in beech forests do not grow in mor (raw humus), a layer of organic matter in various stages of decay containing little or no mineral soil particles. Three possibilities were considered directly or indirectly: (i) tree root competition and litter shading, (ii) phytotoxic and nutrient complexing organic compounds in the soil solution, and (iii) high H-ion concentration in the solution. Rhizomes of eight species (Allium ursinum, Carex sylvatica, Convallaria majalis, Deschampsia flexuosa, Galium odoratum, Poa nemoralis, Stellaria holostea, Stellaria nemorum) were collected in late winter and used in a soil experiment to test reestablishment and growth in untreated mor (soil solution pH 3.6) and after pH was raised to 4.3 by addition of SrCO₃. A flowing solution experiment was used with five of the species mentioned. The solution was composed according to the soil solution of the untreated mor but lacking organic compounds. pH was adjusted to and maintained at 3.6 and 4.3. The experiments showed that even when root competition, litter shading and organic compounds were excluded as limiting factors, only Deschampsia flexuosa, and partly Convallaria majalis, could produce new roots in the untreated mor or in the pH 3.6 solution treatment. Some shoot biomass developed in all species except in Galium odoratum, though significantly less than at pH 4.3.It was concluded that high H-ion concentration of the soil solution precludes establishment and growth of many forest plants in beech forests developed on mor podzols.     

Plantago lanceolata L. and Rumex acetosella L. differ in their utilisation of soil phosphorus fractions

To establish relationships between soil phosphorus (P) fractions and leaf P, a mycorrhizal species (Plantago lanceolata L.) was compared with a typically non-mycorrhizal species (Rumex acetosella L.) in a glasshouse experiment. The plants were grown in 40 soils from non-fertilised, abandoned pastures or abandoned arable fields and leaf P concentration were found to be related to various soil P fractions after six weeks of growth. The differences in the P fractions in soil can account for a large share of the variation in leaf P concentration in both species, but the two species differed in their utilisation of P fractions. Leaf P concentration of R. acetosella was more related to extractable soil P than that of P. lanceolata. Rumex acetosella showed a higher maximum P concentration. The P fractions accounting for the largest share of the variation in leaf P concentration was the Bray 1 extractable and the weak oxalate (1 mM) extractable P, and for P. lanceolata also the Na₂SO₄+NaF extractable P fraction. P extracted with these methods accounted for up to 80% of the variation in P concentration in leaves of R. acetosella and 65% of the variation in leaves of P. lanceolata. More P extractable with weak oxalate, Na₂SO₄+NaF and strong oxalate (50 mM) was released from the soil than was taken up by the plants during the experimental period. The Bray 1 extractable P fraction, however, decreased in both unplanted and planted soils. Phosphatase release was not induced in any of the plants during the experimental period, indicating that they were not mobilising soil organic P. However, some of the methods extracted a large share of the organic P and still explained much of the variation in leaf P concentration. Mycorrhizal colonisation of P. lanceolata was inversely related to the extractable soil P. The consistently fast P uptake of R. acetosella indicates that this species have a high demand for P. The differences in P utilisation between R. acetosella and P. lanceolata could be caused by their different mycorrhizal status.     

Phosphorus enhances Al resistance in Al-resistant Lespedeza bicolor but not in Al-sensitive L. cuneata under relatively high Al stress

Background and Aims

Aluminium (Al) toxicity and phosphorus (P) deficiency often co-exist in acidic soils and limit crop production worldwide. Lespedeza bicolor is a leguminous forage species that grows very well in infertile, acidic soils. The objective of this study was to investigate the effects of Al and P interactions on growth of Lespedeza and the distributions of Al and P in two different Al-resistant species, and to explore whether P can ameliorate the toxic effect of Al in the two species.

Methods

Two species, Lespedeza bicolor and L. cuneata, were grown for 30 d with alternate Al and P treatments in a hydroponics system. Harvested roots were examined using a root-system scanner, and the contents of Al, P and other nutrient elements in the plants were determined using inductively coupled plasma-atomic emission spectroscopy (ICP-AES). Haematoxylin staining was used to observe the distribution of Al in the roots of seedlings. After pre-culture with or without P application, organic acids in the exudates of roots exposed to Al were held in an anion-exchange resin, eluted with 2 m HCl and then analysed using high-performance liquid chromatography (HPLC).

Key Results

Lespedeza bicolor exhibited a stronger Al resistance than did L. cuneata; Al exclusion mechanisms may mainly be responsible for resistance. P application alleviated the toxic effect of Al on root growth in L. bicolor, while no obvious effects were observed in L. cuneata. Much less Al was accumulated in roots of L. bicolor than in L. cuneata after P application, and the P contents in both roots and shoots increased much more for L. bicolor than for L. cuneata. Lespedeza bicolor showed a higher P/Al ratio in roots and shoots than did L. cuneata. P application decreased the Al accumulation in root tips of L. bicolor but not in L. cuneata. The amount of Al-induced organic acid (citrate and malate) exudation from roots pre-cultured with P was much less than from roots without P application; no malate and citrate exudation was detected in L. cuneata.

Conclusions

P enhanced Al resistance in the Al-resistant L. bicolor species but not in the Al-sensitive L. cuneata under relatively high Al stress, although P in L. cuneata might also possess an alleviative potential. Enhancement of Al resistance by P in the resistant species might be associated with its more efficient P accumulation and translocation to shoots and greater Al exclusion from root tips after P application, but not with an increased exudation of organic acids from roots.Key words: Lespedeza bicolor, L. cuneata, Al toxicity, Al resistance, root morphology, phosphorus     

The Monomolecular and Rectangular Hyperbola as Empirical Models of the Response of Photosynthetic Rate to Photon Flux Density, with Applications to Three Veronica Species

The properties of the rectangular hyperbola and monomolecularfunctions, with respect to the photosynthesis/photon flux density(PFD) relationship, are discussed, and the shortcomings of theformer are highlighted. Both models were fitted to data acquiredfrom three closely related Veronica species of contrasting ecology.The non-linear regression algorithms give estimates, with standarderrors, of light saturated photosynthetic rate, light compensationpoint, dark respiration rate, and photochemical efficiency atlow PFD. While the rectangular hyperbola gave almost as gooda fit to the data as the monomolecular for each species, thelight saturated photosynthetic rate estimate given by the formerwas always unacceptably high in comparison with that indicatedby the obvious trend of the data. Moreover, this tendency wasaccentuated if near-saturating PFDs were removed from data sets,and there was a tendency for the fitting algorithm to becomeunstable. No such problems were encountered with the monomolecularfunction, and it is suggested that this be used whenever a simpleempirical model is required to analyze photosynthesis/PFD data. Veronica montana L, Veronica chamaedrys L, Veronica officinalis L, wood speedwell, Germander speedwell, common speedwell, empirical mathematical model, monomolecular function, rectangular hyperbola function, nonlinear regression, photosynthesis, photon flux density, light saturated photosynthetic rate, light compensation point, photochemical efficiency, dark respiration rate     

The Effects of Winter Exposure to Acid Soil Conditions on the Subsequent Survival and Growth of Herbaceous, Forest Perennials: a Preliminary Investigation

For many plant species, growth is limited in acid soils, whichare characterized by high levels of potentially-toxic elementsand low nutrient availability. Although plant-soil interactionsare traditionally studied during the growing season, the highestconcentrations of toxic elements in the soil may occur duringthe winter months. The present study investigated the effectsof a 3-month exposure to either an acid or a reference soil,at temperatures fluctuating around freezing point, on subsequentsurvival and growth of eight herbs (Brachypodium sylvaticum,Carex pilulifera, Geum urbanum, Luzula pilosa, Mycelis muralis,Silene dioica, Stellaria nemorum, Veronica officinalis).Theplants were exposed to ambient weather conditions from Decemberto March, after which they were replanted in fresh referencesoil and transferred to a glasshouse. Their biomass was measured5 weeks later. The plant species differed in their responsesto the soils, in a manner reflecting their natural field distributions.All plants of the most acid-tolerant species survived in bothtreatments, whereas the more sensitive species showed lowersurvival rates after growth in the acid than in the referencesoil. Similar results were found for the regrowth:C. piluliferaandL.pilosa, the most acid-tolerant species, were unaffected by thesoil treatments (ratios between biomass in acid compared toreference soils were 0.8 and 1.1, respectively), whereasG. urbanum,M. muralis, S. nemorumandV. officinaliswere negatively affected(ratios 0.3–0.5). Effects on above- and below-ground biomasswere broadly similar. This preliminary evidence indicates thatsoil chemistry during the winter can be important for both survivaland growth during the vegetative period that follows.Copyright1998 Annals of Botany Company. Acids soils, toxicity, vascular plants, winter exposure.     

Factors Affecting the Restoration of Heathland and Acid Grassland on Pteridium aquilinum–Infested Land across the United Kingdom: A Multisite Study

The variability in the success of Pteridium aquilinum (bracken) control and vegetation restoration has been highlighted as a major issue in the United Kingdom. Experiments were set up at four different regional locations to assess bracken control at the national scale and the impact of restoration practices at the local scale. Bracken control treatments (cutting once or twice per year, a combination of cutting and asulam spraying, and asulam in year 1) were combined with site‐specific treatments designed to restore appropriate heathland or acid grassland vegetation. This article considers the effects on the developing understorey vegetation, testing the following hypotheses: (1) local differences between sites would affect community change; (2) treatments applied to control Pt. aquilinum (same at all sites) influences community change; and (3) treatments applied at the individual site level to restore vegetation influences community change toward the target vegetation. There were a considerable number of spatial effects. It is, therefore, difficult to develop a one‐size‐fits‐all policy for vegetation restoration within a national Pt. aquilinum control strategy. Few bracken control treatment effects were found, and, where they were detected, it was only at single sites. Thus, the development of target vegetation requires a combination of control and restoration treatments that take into consideration the aspects of that site. Only three species, Deschampsia flexuosa, Galium saxatile, and Campylopus introflexus, increased as a direct effect of the control treatments. Vegetation restoration was most successful in the cutting‐twice‐per‐year plots, the treatment with the greatest reduction in Pt. aquilinum cover.     

Genetic adaptation to soil acidification: experimental evidence from four grass species

Anthropogenic acidification has reduced soil pH and released potentially toxic aluminium (Al) ions in many regions. This investigation examines whether increased acidity has caused genetic adaptation to acidic conditions within the grass species Elymus caninus, Poa nemoralis, Deschampsia cespitosa and D. flexuosa. We sampled tussocks (genets) of each species in two regions of southern Sweden, differing in their exposure to acidifying deposition. The tolerance of the genets was tested in a solution experiment with different pH and Al concentrations. Our data suggest that species found at lower pH field locations have a greater tolerance to low pH and high Al levels than species found on less acidic soils. Analysis of variance showed a significant average effect of population and (or) genet in most species; however, we found little evidence of genetic adaptation to acidic conditions at the regional, population and micro-site level. In fact, there was no consistent change in the ranking of populations or genets with varying pH or Al concentration. Based on these results, we hypothesize that phenotypic plasticity rather than genetic adaptation has been favoured as the predominant mechanism to cope with soil acidity in the four grass species.     

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.     

Relationships Between Climate and Flowering of Eight Herbs in a Swedish Deciduous Forest

Effects of annual variation in rainfall, temperature and humidityon flowering abundance of eight temperate woodland plants (Anemonenemorosa, Cardamine bulbifera, Lamiastrum galeobdolon,Oxalisacetosella , Ranunculus ficaria, Stellaria holostea, Viola reichenbachianaand Viola riviniana) were studied during 12 consecutive years(1989–2000) in a hornbeam (Carpinus betulus) forest insoutheast Sweden. Above-average rainfall/humidity in late summerto early autumn of the preceding year increased flowering abundancein L. galeobdolon, O. acetosella, V. reichenbachiana, V. rivinianaand, especially, in R. ficaria, but not in S. holostea and A.nemorosa. Moreover, flowering of R. ficaria and O. acetosellawas positively related to rainfall/humidity during several partsof, or the entire, preceding year. On the contrary, floweringof S. holostea and A. nemorosa was closely related to low valuesof rainfall/humidity in autumn and/or winter of the precedingyear and also to low humidity in the current year in A. nemorosa.Two long periods (3–4 years) of increasing rainfall deficitcoincided with decreasing flowering abundance in most of thespecies, but not with decreasing vegetative development. Temperaturevariability was less consistently related to flowering. A coolperiod during the preceding summer or autumn seemed importantfor flowering in L. galeobdolon, O. acetosella and the Violaspecies, although these relations were, at least partly, causedby interactions with rainfall/humidity. No significant (P <0.05) correlations were found between flowering and the conditionsprevailing in April to May—the main flowering season—ofthe current year. Copyright 2001 Annals of Botany Company Climate, flowering, rainfall, temperature, Anemone nemorosa, Cardamine bulbifera, Lamiastrum galeobdolon, Oxalis acetosella, Ranunculus ficaria, Stellaria holostea, Viola reichenbachiana, Viola riviniana     

Aluminium-induced changes in the profiles of both organic acids and phenolic substances underlie Al tolerance in Rumex acetosa L.

The common sorrel, Rumex acetosa L. is well adapted to acid mineral soils with high availability of phytotoxic Al species. The mechanisms of Al resistance in this species are not established. Our goal was to assess the possible implications of organic acids and phenolic substances in Al detoxification in roots and shoots of this plant. R. acetosa plants were exposed in nutrient solution (pH 4.3) to a non-growth reducing Al concentration of 50 μM Al for 5 days. Exclusion of Al from root tips was visualized by haematoxylin staining. Tissue Al and Ca concentrations were analysed by ICP ES. Root and shoot concentrations of organic acids and phenolic substances were analysed by HPLC. A time-dependent (model II type) Al exclusion pattern in root tips was observed. Nonetheless, high Al concentrations accumulated in roots (1170 μg/g) and shoots (275 μg/g). Aluminium supply enhanced root citrate concentrations but decreased shoot organic acid levels. Aluminium induced high levels of anthraquinone in roots and of catechol, catechin and rutin in shoots. Aluminium resistance in R. acetosa implies both exclusion of Al from root tips and tolerance to high Al tissue concentrations. Citrate in roots and phenolics in shoots may bind Al in non-toxic form. Anthraquinones, as strong antioxidants, may play a role in a general defence response to the root stress.     

Growth,radicle and root hair development ofDeschampsia flexuosa (L.) Trin. seedlings in relation to soil acidity

Deschampsia flexuosa (L.) Trin. is an abundant grass species in the ground flora of acidic beech forests in southern Sweden. Generally, the species is restricted to a limited soil pH range (pH 4–5). The main objective was to study the influence of different soil acidities on germination, initial root development and on the growth of the species. The experiments were carried out under controlled conditions and designed to simulate the physico-chemical conditions present in the field. By using forest soils within the pH range 4.0 to 8.3 and artificial variation in pH (3.2 to 7.6) of soil-water extracts, it was possible to evaluate the influence of soil reaction and the H⁺ per se. In all experiments seeds have been used. Germination was significantly delayed in the very acid soil (pH 4.0) in comparison to the germination in soils within the pH range (4.4 to 6.4). Soil substances, other than the H⁺, might be responsible for this delay in germination, whereas development of the radicle was markedly affected by increasing H⁺ concentrations. Especially the development of root hairs was sensitive to H⁺ and was significantly reduced at a pH<-3.8. By increasing soil acidity the injury symptoms, including curling and discolouring, became more intense and at the highest acidity (pH 3.2) the radicles appeared brown, stunted and the root hairs were lacking. Most favourable growth was obtained at pH 4.4 and 5.0. Soil pH levels above and below this range limited both shoot and root growth. The results showed very good correspondence with observations made in Beech forest soils in southern Sweden, where the species was growing in soils within the pH range 3.9 to 5.1 with a peak growth at pH 4.3. This study shows that in soils at pH≤3.8, the poor development of the radicle may be crucial in the establishment ofDeschampsia flexuosa. Root hair development was more sensitive to soil acidity than radicle elongation. Germination was delayed in very acid Beech forest soils but other factors than the H-ion per se may be responsible for this delay.     

Nitrogen storage forms in nine boreal understorey plant species

Storage forms of N were studied in below-ground structures of nine boreal forest understorey plants. The ericaceous shrubs Vacciniumvitis-idaea and V.myrtillus, the fern Gymnocarpium dryopteris, the grass Deschampsia flexuosa, and the herbs Epilobium angustifolium, Maianthemum bifolium, Solidago virgaurea, Geranium sylvaticum and Trientalis europaea were sampled in early summer and late autumn from plots fertilised with a complete mixture of nutrients and from non-fertilised control plots. Concentrations of total nitrogen, insoluble and soluble proteins, free amino acids and nitrate were measured, and changes in absolute and relative concentrations of these N fractions between early summer and late autumn were used to identify the forms in which the plants store N. In all species studied, the concentration of free amino acids increased both between summer and autumn and in response to fertilisation, while the concentration of protein N increased only in response to fertilisation. Thus, free amino acids appear to have a central role in N storage. In all of the species except G. dryopteris, D. flexuosa and S. virgaurea, arginine dominated the pool of free amino acids and thus arginine was the major form of stored N in most species. In D. flexuosa and S. virgurea, however, asparagine and arginine together were the major forms of stored N, while glutamine was the major free amino acid, and N storage form, in G. dryopteris. Received: 10 March 1996 / Accepted: 22 December 1996     

Effect of Irradiance on Chlorophyll Estimation with the Minolta SPAD-502 Leaf Chlorophyll Meter

Leaf chlorophyll content may be used as an indirect indicatorof crop nitrogen status. Chlorophyll meter values (SPAD values)taken with the Minolta SPAD-502 chlorophyll meter in the shadeplantOxalis acetosellaL. and in winter wheat (Triticum aestivumL.)varied by 15 and 8%, respectively, with variation in irradiance.The lowest SPAD-values were measured at high irradiance. Duringa natural night-day-night cycle SPAD values for winter wheatwere lowest in the middle of the day, highest at low irradianceat dusk and dawn and intermediate in darkness before dawn andafter dusk. The results indicate that irradiance during measurementshould be considered when using the Minolta SPAD-502 chlorophyllmeter for the estimation of crop N-status.Copyright 1998 Annalsof Botany Company Chlorophyll meter, nitrogen, irradiance,Oxalis acetosellaL.,Triticum aestivumL., winter wheat.