Research Article
Is the threshold size for flowering in Cynoglossum officinale fixed or dependent on environment?
- TOM J. DE JONG, LEENTJE GOOSEN-DE ROO, PETER G. L. KLINKHAMER
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- 01 March 1998, pp. 489-496
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In the monocarpic perennial Cynoglossum officinale L. the probability of flowering is related to the size of the plant. In previous work it was observed that this relation varies between years. We hypothesized that variable conditions during the winter, the period of vernalization, explain this variation.
We collected plants from the field in autumn and placed these under different simulated winter conditions in a climate room. In contrast to our hypothesis, the probability of initiating flowering at a given size was not affected by: (a) the temperature during the cold period, (b) the duration of the cold period, or (c) the application of a plant hormone (GA3) or an inhibitor of gibberellin synthesis (paclobutazol) during the cold period. Winter cold is not necessary for floral initiation, and is only required for elongation of the inflorescence. It is unlikely that winter temperature affects the fraction of plants flowering.
Subsequent morphological investigation of flower development in material collected in the field showed that large plants had primordial inflorescences well before vernalization, sometimes as early as August. In plants grown from seeds under constant conditions in a climate room, the probability of initiating the inflorescence differed for plants grown at various temperatures (34·1% at 15°C, 100% at 20°C, and 95% at 25°C). Our results suggest that environmental conditions in August and September, up to 10 months before actual flowering, could affect the fraction of flowering plants.
Foliar application of nitrate or ammonium as sole nitrogen supply in Ricinus communis. I. Carbon and nitrogen uptake and inflows
- ANDREAS D. PEUKE, W. DIETER JESCHKE, KARL-JOSEF DIETZ, LUKAS SCHREIBER, WOLFRAM HARTUNG
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- 01 April 1998, pp. 675-687
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Following a precultivation with pedospheric nitrogen nutrition, nitrate or ammonium solutions were supplied to the shoots of Ricinus plants by spraying (during the experimental period) resulting in an increase of biotic/organic and abiotic/inorganic particles on the surface, which significantly increased wetting of the leaf surfaces. The distribution of particles on the surface of sprayed leaves, in particular crystals around and in stomata, indicated the possible entry of nutrients via thin water films through the stomatal pores in addition to diffusion through the cuticle. Ammonium was taken up more readily than nitrate by the foliage, but both at relatively low rates which caused N limitation. Interestingly, the inorganic N, both in the form of nitrate and even ammonium, was entirely assimilated in the shoots; phloem transport of inorganic N to the root was negligible. The flows of malate, and the acidification of the apoplastic washing solution of leaves in ammonium-sprayed plants pointed to the role of metabolism of malate and excretion of protons in maintaining pH during ammonium assimilation in the shoot. Ammonium-sprayed plants incorporated the N in the same amounts in shoots and roots, only 38% of the shoot-borne N being recycled in the xylem. In nitrate-sprayed plants the root was not only favoured in N partitioning, but even a net export of previously incorporated N from the shoots occurred which reflected the N limitation. The N limitation also affected carbon metabolism, in particular the flows of C, incorporation in the shoot and photosynthesis, which were decreased when compared with data from recent experiments with pedospheric well fed Ricinus. However, there was little difference in C flows between nitrate and ammonium-sprayed plants with respect to respiration, C partitioning and, most interestingly, in relative stimulation of root growth. The loss of C from dark respiration of the shoots was high on a f. wt basis as well as in relative terms, owing to exclusive N assimilation in the shoot. In general the plants invested untargeted increases in root growth as a result of N limitation irrespective of the imposed artificial treatment which made the shoot the site of mineral N uptake.
Induction of genes for the stress proteins PR-10 and PAL in relation to growth, visible injuries and stomatal conductance in birch (Betula pendula) clones exposed to ozone and/or drought
- E. PÄÄKKÖNEN, S. SEPPÄNEN, T. HOLOPAINEN, H. KOKKO, S. KÄRENLAMPI, L. KÄRENLAMPI, J. KANGASJÄRVI
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- 01 February 1998, pp. 295-305
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Well watered (WW) or drought-stressed (DS) saplings of ozone-sensitive and ozone-tolerant (less sensitive) birch (Betula pendula Roth) clones were exposed for 43 d to 0 nl l−1 or 100 nl l−1 ozone. Relative growth rates of leaves, stem, and roots, leaf discolouration, stomatal conductance and induction of genes encoding stress-related proteins PR-10, PAL and a LEA-group protein BP8 were determined. In general, both ozone and drought stress, singly and in combination, increased transcript levels of PR-10 in both clones. This was related to lower induction of PAL (except in older leaves of the tolerant clone), and increased proportions of visibly injured and yellowed leaves in ozone-exposed plants. The clones differed in their stomatal conductance and growth responses. In the less sensitive clone 2, ozone did not affect growth rates, but high stomatal conductance was observed in WW ozone-exposed plants. The more sensitive clone 5 showed, on the contrary, reduced growth rates and low stomatal conductance in WW ozone plants. Interestingly, clone 2 was sensitive to drought stress alone, whereas clone 5 was highly sensitive to ozone and drought stress experienced together. The results show that appearance of visible injuries (necrotic flecks) and enhanced yellowing of leaves coincided with the induction of genes for stress proteins PR-10 and PAL. The short-term growth responses, however, seemed to be separate processes. Additionally, stomatal conductance was related to leaf injuries and growth rates in a complicated manner, emphasizing the complex nature of ozone sensitivity/tolerance mechanisms in birch.
Seasonal changes of leaf surface contamination in beech, oak, and ginkgo in relation to leaf micromorphology and wettability
- C. NEINHUIS, W. BARTHLOTT
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- 01 January 1998, pp. 91-98
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The leaf surfaces of beech, oak and ginkgo have been investigated with respect to contamination with particles during one growing season. Based on the observation that particles are removed from water-repellent leaves by rain (Lotus effect) the three species were selected because they differ in leaf surface micromorphology and wettability. Leaves of beech are smooth, lacked wax crystals and were ±wettable. Those of ginkgo were rough because their cells were convex and were densely covered by wax crystals, resulting in permanent water repellency. Leaves of oak were covered by waxes and were water repellent when young, but, a few weeks after leaf expansion had ceased the waxes were rapidly eroded. These differences in wettability resulted in different amounts of contamination. Ginkgo collected a very small number of particles during the whole vegetation period. In beech the contamination was significantly higher, but fairly constant, whereas oak leaves accumulated particles with age.
Effects of a natural source of very high CO2 concentration on the leaf gas exchange, xylem water potential and stomatal characteristics of plants of Spatiphylum cannifolium and Bauhinia multinervia
- MARÍA DOLORES FERNÁNDEZ, ALEJANDRO PIETERS, CAROLINA DONOSO, WILMER TEZARA, MERCEDES AZKUE, CECILIA HERRERA, ELIZABETH RENGIFO, ANA HERRERA
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- 01 April 1998, pp. 689-697
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The effect of a very high CO2 mole fraction (27000–35000 μmol mol−1) on photosynthesis and water relations was studied during the dry and the rainy season in plants of Spatiphylum cannifolium (Dryand.) Schott and Bauhinia multinervia (H.B.K.) DC. growing near natural cold CO2 springs. Xylem water potential in plants of both species was lowered by drought, high CO2 growth-concentration decreasing it further in S. cannifolium. In plants of both species growing under high CO2 concentration photosynthetic rates measured at a CO2 mole fraction of 1000 μmol mol−1 were higher than in plants growing at ambient CO2 mole fraction and measured at 350 μmol mol−1. The response was the result of a direct effect of CO2 on the photosynthetic machinery. Changes in carboxylation efficiency in response to high CO2 were found during the rainy season, with an increase in S. cannifolium and a decrease in B. multinervia; a significant interaction between growth CO2 concentration and season in B. multinervia resulted from significant effects of both factors. An increase in intrinsic water-use efficiency due to high CO2 was determined in both species by an increase in photosynthetic rate as well as a decrease in leaf conductance. In high-CO2 plants of S. cannifolium a 71% decrease in stomatal density and 73% in stomatal index suggested that CO2 affected stomatal initiation, whereas in B. multinervia an 85% decrease in stomatal index and a 72% decrease in stomatal density indicated that CO2 influenced stomatal initiation as well as epidermal cell expansion. Our results indicate that very high CO2 concentrations did not inhibit photosynthesis in these species, and that growth under high CO2 allowed plants to attain carbon balances higher than those of plants growing under low CO2. This was particularly so during the dry season, since the photosynthetic rates at the corresponding ambient concentration were higher in plants nearer the springs, and carboxylation efficiency and some stomatal characteristics of both species apparently acclimated to high CO2, but patterns were not consistent and bore no obvious relationship to photosynthetic capacity.
Abscisic acid in lichens: variation, water relations and metabolism
- S. DIETZ, W. HARTUNG
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- 01 January 1998, pp. 99-106
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For the first time the well known drought stress hormone abscisic acid, which is involved in regulating processes increasing desiccation tolerance in many plant systems, was analysed in lichens. ABA was detected in all 26 species investigated. In contrast to higher plants and liverworts, the ABA content increased after hydration of air dry lichen thalli and decreased in desiccating lichen material. Experiments with Baeomyces rufus (Huds.) Rebent indicated that the mycobiont might be the major site of ABA biosynthesis. After incubation of hydrated lichen thalli with radioactive ABA for up to 72 h no metabolism to phaseic acid and dihydrophaseic acid could be detected. Fluctuations of internal ABA might be a result of ABA release to the external medium.
Copper–zinc superoxide dismutase is a constituent enzyme of the matrix of peroxisomes in the cotyledons of oilseed plants
- FRANCISCO J. CORPAS, LUISA M. SANDALIO, LUIS A. DEL RÍO, RICHARD N. TRELEASE
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- 01 February 1998, pp. 307-314
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The number and type of isoforms of superoxide dismutase (SOD) and their activities were compared in mitochondria and peroxisomes isolated from cotyledons of three different oilseed seedlings. Mitochondrial and peroxisomal isoforms of SOD could be distinguished in nondenaturing polyacrylamide gels by their differential sensitivities to KCN and/or H2O2. The type of SOD was not the same for each organelle in each of the three oilseed species. For example, a single Mn–SOD was found in cotton and cucumber mitochondria, whereas four CuZn–SODs were present in mitochondria from sunflower. At least one CuZn–SOD isoform was found in the peroxisomes of all three species. Cucumber peroxisomes contained both a CuZn–SOD and a Mn–SOD, cotton peroxisomes contained a single CuZn–SOD, whilst four separate CuZn–SODs, but no Mn–SOD were found in sunflower peroxisomes. Using antibodies against CuZn–SOD from watermelon peroxisomes or from chloroplasts of Equisetum, a single polypeptide of c. 16·5 kDa was detected on immunoblots of peroxisomal fractions from the three species. Post-embedment, electron-microscopic double immunogold-labelling showed that CuZn–SOD, with malate synthase used as marker enzyme of peroxisomes, was localized in the matrix of these organelles of all three species. These results suggest that CuZn–SOD is a characteristic matrix enzyme of peroxisomes in oilseed cotyledons.
The long-term epidemic of Claviceps purpurea on Spartina anglica in Poole Harbour: pattern of infection, effects on seed production and the role of Fusarium heterosporum
- A. F. RAYBOULD, A. J. GRAY, R. T. CLARKE
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- 01 March 1998, pp. 497-505
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An epidemic of the ergot fungus (Claviceps purpurea (Fr.) Tul), a non-systemic floral pathogen, appeared in populations of Spartina anglica C. E. Hubb in Poole Harbour in the 1980s. Over 70% of inflorescences were infected between 1985 and 1995. Between 1983 and 1995, there was no consistent pattern in the rank order of sites with respect to the proportion of inflorescences infected, and variation in the amount of infection was not related to inflorescence density. Among years, there was significant variation in the total biomass of ergot per inflorescence, but no significant difference in the number of ergots per inflorescence. Log–log regressions of total weight of ergot per inflorescence against the number of ergots per inflorescence showed that in all years resources for each additional ergot were limited.
There was no statistically significant difference between the number of seed set on infected and uninfected inflorescences in 1985 or 1995. Further analysis showed that, compared with uninfected inflorescences, there was higher seed output from inflorescences with fewer than 10% of spikelets infected, which balanced reduced seed output from heavily infected inflorescences. At high levels of ergot infection, a lower proportion of uninfected spikelets set seed, compared with spikelets on uninfected inflorescences. This suggests that conditions which favour ergot growth are detrimental to seed production. Compared with uninfected inflorescences, mean and total seed weights were significantly lower in inflorescences with >10% of spikelets infected.
The fungus Fusarium heterosporum Nees ex Fr. was found in association with the ergots on Spartina. There was a significant positive correlation between the average severity of Fusarium infection and the number of ergots on an inflorescence. There was a non-significant negative correlation between severity of Fusarium and mean ergot weight, when ergot number per inflorescence was held constant. These data suggest that any negative effect of Fusarium on Claviceps is small.
The high, and temporally and spatially uniform levels of ergot infection probably result from genetic uniformity of Spartina and the lack of zonation in the salt marshes of Poole Harbour. This situation is likely to persist for the foreseeable future.
Ecotypic adaptation in Austrocedrus chilensis in cuticular hydrocarbon composition
- RICHARD S. DODD, ZARA A. RAFII, ARIEL B. POWER
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- 01 April 1998, pp. 699-708
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An homologous series of hydrocarbons ranging from C21 to C37 (was detected in foliar cuticular waxes of Austrocedrus chilensis (D. Don) Pic.-Ser. & Bizz., by gas chromatography. Qualitatively, the chromatograms were similar for all individuals sampled from 29 natural populations in Chile and Argentina, showing C33 and C35 alkanes to be dominant. Quantitatively, the chromatograms varied significantly among populations. Multivariate analyses showed that arid zone populations from mediterranean Chile and the Patagonian steppe were different from one another and were distinct from mesic populations close to the Valdivian rainforest vegetation zone. Mediterranean Chilean populations were characterized by higher concentrations of the longer carbon-chain alkanes, but retained equal amounts of shorter-chain homologues as the mesic populations. A greater variation in chain lengths and higher concentrations of longer carbon chains in the mediterranean populations is consistent with a model for the adaptation to reduced cuticular permeability. Partial Mantel matrix tests revealed significant climatic and inter-population distance effects with taxonomic distances based on multivariate and univariate hydrocarbon data. Annual rainfall was overall the most significant factor, particularly in regressions with the shorter-chain hydrocarbons. Annual mean temperature was most significant for the longer-chain hydrocarbons. This suggests, on the one hand, ecogenic adaptation to both temperature and precipitation of cuticular hydrocarbon composition, and, on the other hand, a weaker, but important effect of gene flow in determining hydrocarbon composition in this species. The northernmost populations at San Felipe and San Gabriel were the most distinctive. This could result from the effects of random changes in allele frequencies and/or to founder effects in isolated and small populations. These populations show some East-Andean affinities presumably due to historic migration patterns.
Fungal infections of the phytoplankton: seasonality, minimal host density, and specificity in a mesotrophic lake
- HARALD HOLFELD
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- 01 March 1998, pp. 507-517
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Phytoplankton infections by fungal parasites in the upper, mixed layer of a mesotrophic northern temperate lake were analysed according to the following parameters: host and parasite species, host population density and prevalence of infection, resting spore formation by the parasite, and the lowest host density at which parasites appeared. The phytoplankton taxa recorded included the Cyanobacteria, Dinomastigota, Chrysophyceae, Bacillariophyceae, Chlorophyceae, Cryptophyceae and Haptophyceae, but infection was never found in the last two classes. The parasites belonged almost exclusively to the monocentric Chytridiomycetes. Fungal epidemics occurred at all times of the year. Parasites appeared at population densities as low as about 1 cell ml−1 in some host species, with infection prevalence sometimes exceeding 80%. The proportion of the total phytoplankton biovolume infected by fungi was usually much <1%, but occasionally reached 10%. Parasitism proved to be highly species-specific, with one parasite species usually infecting only one host species. In the case of Zygorhizidium planktonicum, which infected both Asterionella formosa and Synedra acus, there is evidence that two species-specific formae speciales, each infecting only one of these two host species, are present in the lake.
Elevated carbon dioxide ameliorates the effects of ozone on photosynthesis and growth: species respond similarly regardless of photosynthetic pathway or plant functional group
- JOHN C. VOLIN, PETER B. REICH, THOMAS J. GIVNISH
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- 01 February 1998, pp. 315-325
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Due to their different physiological effects, elevated carbon dioxide and elevated ozone might have interactive impacts on plants, and differentially so on plants differing in photosynthetic pathway and growth rate. To test several hypotheses related to these issues, we examined the physiological, morphological and growth responses of six perennial species grown at various atmospheric concentrations of carbon dioxide and ozone. The species involved (two C3 trees: Populus tremuloides Michx., Quercus rubra L.; two C3 grasses: Agropyron smithii Rybd., Koeleria cristata L.; two C4 grasses: Bouteloua curtipendula Michx., Schizachyrium scoparium Michx.) differed in growth form, stomatal conductance and photosynthetic pathway. In situ photosynthesis, relative growth rate (RGR) and its determinants (leaf area ratio, specific leaf area, leaf weight ratio and root weight ratio) were determined via sequential harvests of seedlings that were grown in all combinations of 366 or 672 μmol mol−1 CO2 and 3 or 95 nmol mol−1 O3 over a 101-d period. Elevated CO2 had minimal effect on either photosynthesis or RGR. By contrast, RGR for all six species was lower in high O3 concentrations at ambient CO2, significantly so in A. smithii and P. tremuloides. Five of the six species also exhibited reductions in in situ photosynthesis at ambient CO2 in high-O3-grown compared with low-O3-grown plants. For all species, these O3-induced reductions in RGR and photosynthesis were absent in the elevated CO2 environment. Root weight ratio was significantly reduced by elevated O3 in A. smithii and P. tremuloides in ambient but not elevated CO2. Species with high stomatal conductance were the most susceptible to oxidant injury, while those with low stomatal conductance, such as the C4 species and Q. rubra, were not as detrimentally affected by O3. Elevated levels of CO2 will reduce stomatal conductance and O3 uptake, and might therefore reduce the potential for oxidant damage. However, there was a stronger relationship of the percent reduction in whole-plant mass due to O3, related to the ratio of photosynthesis to stomatal conductance. In general, results of this study of six functionally diverse plant species suggest that O3 pollution effects on carbon balance and growth are likely to be ameliorated by elevated concentrations of atmospheric CO2.
Suppression of hyphal growth of soil-borne fungi by dune soils from vigorous and declining stands of Ammophila arenaria
- W. DE BOER, P. J. A. KLEIN GUNNEWIEK, J. W. WOLDENDORP
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- 01 January 1998, pp. 107-116
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A study was carried out to determine whether expansion of marram-grass stands (Ammophila arenaria (L.) Link) on acidic inner Dutch coastal dunes was caused by suppressiveness of soils from these stands against three potential pathogenic fungi of marram grass, namely Fusarium culmorum (W. G. Sm.) Sacc., Phoma exigua Desm. and a Ulocladium sp. The suppressiveness of the acidic inner dune soils was compared with that of lime-rich dune soils from vigorous and declining marram-grass stands. Suppressiveness of the dune soils against the saprotrophic fungi Chaetomium globosum Kunze: Fr, Mucor hiemalis Wehmer and Trichoderma harzianum Rifai was also determined. All fungi had been isolated from marram-grass roots. Suppressiveness was determined by comparing the formation of hyphae from potato–dextrose agar discs into (layer method) or on top of (surface method) dune soils with that of controls consisting of sterile, acid-washed beach sand. The growth of the three root-infecting fungi was strongly inhibited in all soils regardless of the method used. Hence, there were no indications that the potential pathogenic fungi were selectively suppressed by the acidic dune soils and, consequently, the results did not give any indication for the involvement of a fungal component in the decline of marram grass. Growth of the saprotrophs C. globosum and M. hiemalis was much less inhibited than that of the root-infecting fungi. Growth of T. harzianum was strongly inhibited in alkaline soils but not in the acid ones. The suppression of fungal growth could be partly or completely eliminated by a microwave treatment, indicating that biological components of the soil were essential to suppressiveness. The suppression of the fungi by colonies of dune soil micro-organisms on water-agar differed considerably from soil alone. Yet, all methods indicated the occurrence of general suppressiveness against fungi by dune soils, irrespective of the origin of the soil samples. This suppressiveness was probably not due to direct competition with other soil micro-organisms for nutrients but to inhibiting compounds produced by the soil micro-organisms.
Developmental and morphological responses of mycelial systems of Stropharia caerulea and Phanerochaete velutina to soil nutrient enrichment
- DAMIAN P. DONNELLY, LYNNE BODDY
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- 01 March 1998, pp. 519-531
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Patterns produced by foraging mycelial systems of decomposer fungi might be the visual evidence of strategies for obtaining differently distributed carbon sources. Mycelial systems might also be responsive to unevenly distributed mineral nutrients in the soil. The nature of the mycelial response might differ between species inhabiting eutrophicated disturbed sites, such as Stropharia caerulea (Kriesel), and inhabitants of more typical woodland soils, such as Phanerochaete velutina DC.: (Pers.) Parmasto. We examined the effect of enrichment of soil by addition of different concentrations of N, P and potassium on inoculum decay rate and on extension rate, biomass production and fractal dimension (determined by image analysis) of extra-resource mycelia of S. caerulea and P. velutina extending from wood blocks across compacted non-sterile soil in trays.
S. caerulea was more responsive than P. velutina to both type and quantity of soil nutrient enrichment. P. velutina grew more rapidly, produced more biomass and decayed inocula more rapidly than S. caerulea under most conditions, rates of growth and resource decay of the former remaining stable over a range of soil nutrient enrichments. In contrast, morphology of S. caerulea altered on enriched soils compared to controls with greater and more pronounced fanning (divergence of separate hyphae) of the foraging mycelial front. These changes were reflected in fractal values. Extension rate and biomass production of S. caerulea was less than controls on N-amended soil, but greater on P-amended soils, with the greatest biomass produced on 0·13–0·27 mg P g−1 air-dried soil. Decay rate of inocula was often positively related with mycelial production rates. These responses indicate extra-resource mycelium of S. caerulea is able to respond rapidly to a range of soil nutrient enrichments, and this is discussed in terms of the different foraging strategies exhibited by the two fungi.
The response of Norway spruce seedlings to simulated acid mist
- LUCY J. SHEPPARD, IAN D. LEITH, MAUREEN B. MURRAY, J. NEIL CAPE, VALERIE H. KENNEDY
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- 01 April 1998, pp. 709-723
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Four pot experiments are reported in which Norway spruce (Picea abies (L.) Karst) seedlings, of different nutrient status, were treated with acid mist for one growing season in open-top chambers (OTCs). Combinations of H+, SO42−, NH4+ and NO3− were applied at different frequencies of application and supplying different doses of S and N kg ha−1. Plant growth, visible injury, frost hardiness and nutrient status were observed. These experiments were undertaken to improve our understanding of the interaction of environmental factors such as nutrition and mist-exposure frequency on seedling response to N and S deposition.
Both acidity ([les ]pH 2·7) and SO42− ions were necessary to induce visible injury. Mist containing SO42−, H+ and to a lesser extent NH4+ significantly reduced winter frost hardiness. Increasing the misting frequency, and to a lesser extent the overall dose, increased the likelihood of acid mist causing visible injury and reducing frost hardiness. Post-planting stress, low N status and needle juvenility increased the likelihood of acid mist causing visible injury. Increased plant vitality, adequate N status and growth rate reduced the likelihood of acid-mist-induced reductions in frost hardiness.
Principles underlying the responses of spruce seedlings treated in controlled conditions to acid mist are discussed.
The arbuscular mycorrhizal fungi of Hyacinthoides non-scripta I. Diversity of fungal taxa
- JAMES MERRYWEATHER, ALASTAIR FITTER
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- 01 January 1998, pp. 117-129
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The arbuscular mycorrhizas of bluebell (Hyacinthoides non-scripta (L.) Chouard ex Rothm.) involve several symbiotic fungi of the order Glomales. We have previously simplified the system by ignoring the taxonomic diversity of the fungi, but it is unlikely that all fungal species contribute in the same manner or to the same extent to the functioning of the symbiosis. To discover how many and which fungi take part in the bluebell mycorrhiza we sought to identify the range of arbuscular mycorrhizal (AM) fungi found in bluebell roots sampled during a complete growing season, September to June.
Although the taxonomy of the Glomales by their spores is not yet fully understood, identification is, to a large extent, possible. Arbuscular mycorrhizal communities are usually characterized by their spores but, since spores can rarely be directly associated with individual plants or plant species, a more satisfactory approach would be to identify fungal symbionts where they interact with the host plant, in the roots. Unfortunately, the intraradical mycelia of the fungi are less easily distinguished than the spores and, as yet, classification is possible only to the family level.
We have developed a method whereby different AM fungal taxa in the roots of bluebell can be distinguished by objective assessment. A large suite of morphological characteristics of the fungi in roots was recorded in samples taken at monthly intervals. Hierarchical cluster analysis of the resulting data separated six distinct AM fungal morphotypes (Scutellospora type, Acaulospora type, three Glomus types and fine endophytes) and a classification system created by which identification by eye was possible.
We compared the fungi in roots with glomalean spores in soil from the root zone of the same bluebell plants. Two species occurred in most samples, Scutellospora dipurpurescens Morton & Koske (emend. Walker, 1993) and Acaulospora koskei Blaskowski. A further six occurred sporadically, five Acaulospora spp. and Glomus rubiforme Gerdemann & Trappe) Almeida & Schenck (=Sclerocystis rubiformis). The presence of a single species of Scutellospora was consistent with a Scutellospora root morphotype which varied little. By contrast, the diversity of Acaulospora in the spore assemblage was reflected by variation within the Acaulospora morphotype. Glomus spores were very rarely found in field collections, yet Glomus morphotypes were found to be an important component of the bluebell mycorrhiza.
Because some important species are not represented in spore assemblages in the field and those that are found can only be associated with vegetational groups, not individual plants or single species, glomalean spore populations provide only a partial account of the fungi which contribute to arbuscular mycorrhizas. Although it is still not possible to identify AM fungi in roots with the same precision as their spores, the method reported here permits assessment of diversity in the roots of individual plant species, which may be applied to the investigation of mycorrhizal function and demography in natural ecosystems.
Daily embolism and refilling of xylem vessels in the roots of field-grown maize
- M. E. McCULLY, C. X. HUANG, L. E. C. LING
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- 01 February 1998, pp. 327-342
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Embolisms in the vessels of maize axile roots of different types were observed directly after rapid freezing of intact, functioning roots in the field, by cryo-scanning electron microscopy. Quantification of the degree of embolization in each root was made by counting empty and full vessels of both the late and early metaxylem (LMX & EMX), and expressed as percent embolized vessels of the LMX, and %EMX poles containing embolized vessels. Contents of the connecting xylem (CX) at branch root junctions, and of xylem in branch roots were observed also, but not systematically quantified. Records of % embolized vessels were made from dawn to dusk on summer days in Ottawa under moderate irradiance, and in Canberra under high irradiance. Measurements in Canberra were supported by estimates of irradiance, of stomatal conductance, and of chamber balance pressure of bagged and unbagged leaves. Soon after sunrise embolisms appeared in all types of vessel, at balance pressures c. 300–400 kPa, and increased rapidly with increasing irradiance. During the middle of the day % embolized vessels reached a maximum (LMX ≈70% in Ottawa, and ≈80% in Canberra). At all times the EMX vessels were less embolized. The midday maximum was brief in Ottawa, and % embolized vessels fell to a low value during the afternoon. In Canberra the maximum was prolonged into late afternoon. By dusk nearly all vessels were once again filled with sap. The balance pressures measured during vessel refilling in Canberra ranged from 500 kPa to 1200 kPa. At all times of the day sap was seen entering some embolized vessels. Almost all were refilling by mid- to late-afternoon. Such refilling was especially frequent at junctions of branch roots with the axile roots. X-ray microanalysis of the sap entering the vessels, and of the liquid filling or partly filling vessels, showed the concentration of mineral solutes present in the sap was below the threshold of detection (≈12 mM). These results are discussed in relation to current opinions about embolisms and vessel refilling.
Cluster root development in Grevillea robusta (Proteaceae). I. Xylem, pericycle, cortex, and epidermis development in a determinate root
- KEITH R. SKENE, JOHN A. RAVEN, JANET I. SPRENT
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- 01 April 1998, pp. 725-732
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The cluster roots of Grevillea robusta A. Cunn. ex R. Br. are composed of determinate rootlets that stop growing, but remain physiologically active for several months. Their apical organization, both before and after maturation, was studied by light and transmission electron microscopy. Each cell layer forms a dome, with an initial cell at its end. Xylem elements form a complicated triarch array at the base of the rootlet, passing along the rootlet as two files, and then joining at the tip to form a single file, surrounded by six pericycle cells. At the base of the rootlet, shorter xylem cells and thick-walled support cells are visible. A root cap, present in rootlets grown in vermiculite, was eventually displaced by root hair growth. Rootlets grown in Hoagland's solution lacked root caps and were significantly shorter than those grown in vermiculite. Cell fate was analysed in terms of cell position and is discussed in terms of pattern and development.
The arbuscular mycorrhizal fungi of Hyacinthoides non-scripta II. Seasonal and spatial patterns of fungal populations
- JAMES MERRYWEATHER, ALASTAIR FITTER
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- 01 January 1998, pp. 131-142
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Roots of bluebell (Hyacinthoides non-scripta (L.) Chouard ex Rothm.) are colonized by a range of fungal symbionts from several genera of the order Glomales. Using the identification scheme described in Merryweather & Fitter (1998), arbuscular mycorrhizal (AM) fungi in bluebell roots were quantified throughout the single growing season of 1994–5 and compared with populations of spores found in the soil around the roots.
In the early part of the growing season, when its activity is entirely subterranean (autumn and winter), bluebell habitually associates with a Scutellospora morphotype which is almost certainly S. dipurpurescens Morton & Koske (emend. Walker, 1993) whose spores occur in the root zone. This is the time of maximum phosphorus inflow and bulb-stored carbohydrate utilization by this mycorrhiza. A diverse flora of other AM fungal morphotypes (Acaulospora and Glomus), which might also form mycorrhizas with other plants in the vicinity of bluebells, invade the roots later in the season (spring), when P inflow is reduced and carbohydrate is available as fresh photosynthate. Their contribution to the mycorrhiza might be less than that of Scutellospora, particularly in terms of P assimilation.
Both AM fungi in roots and glomalean spores recovered from soil around bluebell roots showed a significant degree of correlation with the vegetation within which the test plants grew. In the case of AM fungi in roots, Scutellospora showed no special preference for either, but Glomus correlated with a canopy of sycamore (Acer pseudoplatanus L.) and Acaulospora with oak (Quercus petraea (Mattuschka) Liebl.). Spores which most closely resembled S. dipurpurescens and Acaulospora gerdemannii Schenck & Nicolson were significantly more numerous under sycamore, but a spore like Acaulospora koskei Blaskowski, the most numerous and frequently encountered glomalean spore in the system, showed no preference for areas dominated by either tree. There was no significant relationship between AM fungal populations in bluebell roots and glomalean spores recovered from associated soil.
The two spore taxa most frequently found in the vicinity of bluebell roots (A. koskei and S. dipurpurescens) were also found in lower numbers in soil from a region of the field site in which bluebell was absent, indicating that the main bluebell AM fungi do not exclusively associate with that host.
Genetic structure of a natural population of the ectomycorrhizal fungus Suillus pungens
- PIERLUIGI BONELLO, THOMAS D. BRUNS, MONIQUE GARDES
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- 01 March 1998, pp. 533-542
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Sequence-based markers were developed to study the genetic structure and reproductive biology of the ectomycorrhizal fungus Suillus pungens Thiers & Smith in a Bishop pine (Pinus muricata D. Don) forest. Six different basidiome genotypes were found in a 1200 m2 area. Five of the six genotypes were represented by single basidiomes. The remaining genotype comprised 13 basidiomes and covered an area of at least 300 m2, with maximum measured dimensions of 40 m and 14 m. This is the largest genet of an ectomycorrhizal fungus described to date, and is likely the result of vegetative growth, because analysis of single spore isolates eliminates the possibility of genetic identity resulting from either apomixis or fortuitously indistinguishable recombinant genotypes. Genetic analysis also shows that although out-crossing appears to predominate in the population, at least a low percentage (1·4%) of spores are secondarily homothallic.
The combination of extensive vegetative growth and abundant fruiting suggests S. pungens utilizes more carbon than might be expected for a species which accounts for <3% of the total ectomycorrhizal abundance at the site. Additional carbon might come from either more efficient host–fungus transfer, pooling of carbon from the roots of different host plants, or saprophytism.
Carbohydrate composition and invertase activity of the leafy liverwort Porella platyphylla
- MARIANN MARSCHALL, MICHAEL C. F. PROCTOR, NICHOLAS SMIRNOFF
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- 01 February 1998, pp. 343-353
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The major soluble carbohydrates in the desiccation-tolerant leafy liverwort Porella platyphylla (L.) Lindb. are sucrose and a homologous series of fructans including the trisaccharide 1-kestose. Exogenous glucose and fructose (10 mol m−3) did not affect the composition of the soluble carbohydrate pool. Sucrose caused an increase in the fructan pool. Sucrose also inhibited photosynthetic oxygen evolution and respiration. The fructan pool was maintained in preference to sucrose during dark starvation. Low temperature and low water potential increased the fructan pool whereas desiccation increased the proportion of high molecular weight fructan. Acid invertase activity was detected in a taxonomically diverse range of liverworts but was very low or undetectable in a range of mosses. The invertase activity from P. platyphylla was partially purified by ammonium sulphate precipitation. The reaction products of the partially purified enzyme were equimolar glucose and fructose. Kestose and higher DP fructans were not detected suggesting that, at least under the assay conditions used, the enzyme does not have sucrose[ratio ]sucrose fructosyl transferase activity. The pH optimum was 4·5–5 and the Km for sucrose was 1·7 mol m−3. Pyridoxal hydrochloride (5 mol m−3) caused 50% inhibition. The coexistence of sucrose and invertase suggests that either the invertase is inactive in vivo or is in a different subcellular compartment from sucrose. The pH response shows that it would have very low activity at cytosolic pH. A large acidic vacuole was detected in P. platyphylla leaf cells by neutral-red staining in which either invertase or sucrose could be sequestered. Rehydrating desiccated P. platyphylla for 10 min resulted in a 60% loss of extractable invertase activity. By contrast, extractable malate dehydrogenase activity increased during rehydration. Rehydrating desiccated leaves caused an increase in glucose and fructose suggesting that the sucrose pool was susceptible to invertase at this time. It is suggested that the partial inactivation of invertase during rehydration minimizes sucrose hydrolysis while membrane structure and subcellular compartmentation are re-established.