Carbohydrate Metabolism and Frost Hardiness in Pine and Spruce Seedlings Grown at Different Photoperiods and Thermoperiods

Large changes occur in carbohydrate contents of pine (Pinus silvestris L.) and spruce (Picea abies (L.) Karst.) seedlings cold-hardened by photoperiod or by combined photo- and thermo-period. The largest change is in sucrose content, which is almost doubled after six weeks short-day (6/18 h) treatment; and more than doubled (spruce) or more than tripled (pine), when also temperature is lowered (10/5°C). Development of frost hardiness is strongly correlated with the change in carbohydrate contents. At dehardening, the carbohydrate content decreases rapidly, especially in pine, and the raffinose formed during the rest period disappears within 2–4 weeks. Frost hardiness decreases in parallel. The content of soluble carbohydrates may thus play a role in frost hardiness, although it is not the only factor. Bud formation at cold acclimation is not directly correlated with the changes in carbohydrate content and hardiness.     

Frost tolerance and biochemical changes during hardening and dehardening in contrasting white clover populations

Successful winter survival of perennial plants, like white clover, is dependent on proper timing of both hardening and dehardening. The purpose of this study was to investigate the regulation of these processes in two cultivars (AberCrest and AberHerald) and two Norwegian ecotypes (Særheim collected at 58°46′N lat. and Bodø at 67°20′N lat.) of white clover (Trifolium repens L.). For hardening and dehardening, plants were exposed to controlled temperature conditions and frost hardiness of stolons was tested by programmed freezing at the rate of 3°C per hour. In addition, stolons were analysed for starch, soluble sugars and soluble amino acids. Cultivars AberCrest and AberHerald, selected for growth at low temperature and winter hardiness in the United Kingdom, were significantly less hardy than the Norwegian populations. After six weeks of hardening (2 weeks at 6°C and 4 weeks at 0.5°C), estimated LT₅₀ values were ?13.8, ?13.0, ?17.8 and ?20.3°C for AberCrest, AberHerald, Saerheim and Bodø, respectively. The rate of dehardening increased with increasing temperature. At low temperature (6°C), the northern ecotype from Bodø was more resistant to dehardening than AberHerald. However, at 18°C the absolute rate of dehardening (°C day^?1) was twice as high in Bodø as in AberHerald plants. Stolon elongation during dehardening was initiated at lower temperatures in AberHerald than in plants of the Bodø ecotype. The content of total soluble sugars, sucrose and the amino acids proline and arginine were significantly higher in hardy plants of Bodø than in those of AberHerald. Sucrose levels decreased during dehardening and correlations between sucrose content and LT₅₀ during this process were statistically highly significant for both Bodø and AberHerald. The least hardy populations of white clover were characterized by thick stolons, long internodes and large leaves.     

Relationship between frost tolerance and sugar concentration of various bryophytes in summer and winter

Summary Frost resistance, measured via the photosynthetic capacity after freeze-thaw treatment, and concentrations of sucrose, glucose and fructose of thalli of seven species of Bryidae and one species of Marchantiidae were determined from January to March and June to September, respectively. A distinct increase in cold tolerance from summer to winter was found in Polytrichum formosum Hedw., Atrichum undulatum (Hedw.) P. Beauv., Plagiomnium undulatum (Hedw.) Kop., Plagiomnium affine (Funck) Kop., Mnium hornum Hedw. and Pellia epiphylla (L.) Corda. While the frost resistance of the musci differed in summer and winter by 15° to more than 25° C, the hardening capacity of the thalloid liverwort was comparably low. Except in Mnium hornum, the increase in frost hardiness was accompanied by rise of the sucrose concentration in the cells, but insignificant changes in glucose and fructose contents. In contrast, Brachythecium rutabulum (Hedw.) B.S.G. and Hypnum cupressiforme Hedw. already exhibited high frost tolerances in summer, which coincided with high sucrose levels in the tissue, comparable to those found in other musci during the winter. Highly frost-resistant musci had total sugar concentrations around 90–140 mM, of which at least 80% and often more than 90% was sucrose. Artificial degradation of sucrose during exposure of mosses to higher temperatures resulted in a decline in cold hardiness. The results signify that the concentration of sugars, mainly of sucrose, may be important for the frost tolerance of bryophytes.     

Changes in cold hardiness of introduced and native interior Alaskan evergreens in relation to water and lipid content during spring dehardening

Needle hardiness of introduced yellow pine, Pinus banksiana Lamb., lodgepole pine, P. contorta Dougl, and native white spruce, Picea glauca (Moench) Voss, were assessed by the effective prefreezing temperature method. Yellow pine needles were less hardy than lodgepole pine or white spruce needles in Alaska on each date measured. Although hardiness decreased in springtime in all species, decreases in hardiness in yellow pine began before temperatures were above ?20°C, apparently in response to day length, while decreases in hardiness in lodgepole pine and white spruce began only when mean temperatures were above 0°C. Hardiness was increased by decreasing the water content of yellow pine and spruce needles. However, only the latter increased its field hardiness by decreased water contents, and only to a small degree. Large decreases in phospholipid occurred during the dehardening period, indicating the presence of major membrane-associated changes. However, changes in hardiness did not closely parallel those in phospholipid; hardiness decreased before phospholipid did in spruce and after phospholipid did in lodgepole pine. In yellow pine, changes in hardiness were more closely related to changes in phospholipid content. Decreases in phospholipid appeared to be correlated with the day length in all species.     

Oxygen Diffusion from the Roots of Woody Species

Application of the‘polarographic’ technique for studying oxygen diffusion from roots has provided preliminary results which establish that oxygen passes through the woody species Salix atrocinerea Brot., Salix fragilis L., Salix repens L., and Myrica gale L., in the gaseous phase as it does in other wetland species. Entry into the shoots occurs through the bark directly above the water table and in the willow cuttings the effective length of shoot for gas intake was the basal three centimetres above the water table. The length of shoot involved was longer in Myrica gale and the roots were of the normal (un-nodulated) type. Trials on Alnus glutinosa (L.) Gaertn. provided no conclusive results and it is thought that this was perhaps because measurements were on nodulated roots only.     

Quantitative Analysis of Cold Hardening and Dehardening in Lolium

The change in cold hardiness of three Lolium multiflorum Lam.varieties was followed in plants exposed to hardening or dehardeningconditions at a range of temperatures. Hardening and dehardeningwere analysed as quantitative processes dependent upon temperatureand time. Their time courses changed exponentially to an asymptotewhich was logistically related to temperature. Both componentsof the model were fitted simultaneously. Parameters of biologicalinterest, such as the initial rates of hardening and dehardeningfor a given temperature and the percentage of the process completedin a given time, were derived and compared for varieties ofcontrasting hardiness. The analysis demonstrated the importanceof dehardening in determining hardiness. Similar results wereobtained when the model was applied to hardening and dehardeningdata for Lolium perenne L. The potential of this quantitative analysis for distinguishingbetween possible mechanisms of cold hardiness is discussed andfurther experiments required to characterise the kinetics ofhardening and dehardening more fully are identified. Lolium multiflorum, Lolium perenne, cold hardening, cold dehardening, processes, model, varieties     

Soil water status influences plant nitrogen use: a case study

We studied differences in nitrogen use efficiency (NUE) among six species [Calamagrostis epigejos (L.) Roth., Carex duriuscula C.A. Mey., Phragmites communis (L.) Trin., Salix gordejevii Y.L. Chang, Salix cheilophila Schneid., and Typha minima Funk.] growing in two contrasting habitat types, i.e., a riverine wetland with high water supply and a riparian zone with low water supply. The two sites were different in soil water supply, but not in nitrogen supply. Here, NUE was defined as the total net primary production per unit nitrogen absorbed. There was no significant difference in NUE between the species growing in the riverine wetland (Carex duriuscula, P. communis, S. cheilophila, T. minima) and the species growing in the river bank (Carex duriuscula, Calamagrostis epigejos, P. communis, S. gordejevii). We further analyzed NUE as the product of the nitrogen productivity (A, the rate of dry matter production per unit of nitrogen in the plant) and the mean residence time of nitrogen (MRT, the period of time a unit of nitrogen is present in the plant). The species growing in the riverine wetland had larger A but lower MRT than the species growing in the river bank. There was an inverse relationship between A and MRT. Consequently, NUE was similar among species and habitats. These results suggested that environmental factors, such as soil water supply, can influence N use by plants.     

Chemical composition studies on some aquatic macrophytes in three Scottish lochs. I. Chlorophyll,ash, carbon,nitrogen and phosphorus.

The chlorophyll, ash, carbon, nitrogen and phosphorus levels in seven species of freshwater macrophytes (Juncus effusus L., Iris pseudacorus L., Carex rostrata Stokes, Glyceria maxima (Hartm.) Holmberg, Nuphar lutea (L) Sm., Polygonum amphibium L. and Schoenoplectus lacustris (L) Pallas) in three Scottish lochs of different trophic levels were studied during 1975. Mean chlorophyll levels varied from a minimum of 1.73 mg g^–1 dry weight in Balgavies Loch Juncus to 10.22 mg g^–1 dry weight for Forfar Loch Iris. Carbon contents ranged from 450 to 520 mg g^–1 ash-free dry weight. For ash, nitrogen and phosphorus, significant differences in mean concentrations were detected among plant species as well as within one plant species growing in different lochs. Positive correlations were apparent between the degree of eutrophication in the study areas and the amount of ash, phosphorus and nitrogen present in the plants growing in them.     

Photoperiod control of the initial phase of frost hardiness development in Pinus radiata

Abstract. The development of frost hardiness in Pinus radiata was investigated to establish whether there is quantitative relationship between photoperiod and the hardening process. Three controlled environment experiments were carried out. In the first, seedlings were exposed to a photoperiod that shortened from 13 h at a rate of 3 min d^?1 to 9.5 h. At intervals, the photoperiod was either held constant or lengthened. In the second experiment, seedlings were exposed to one of five constant photoperiods between 9 and 12 h for up to 90 d. In the third, seedlings were exposed to photoperiods shortening at rates of 1 or 5 min d^?1. Frost hardiness was also measured during the natural photoperiod-controlled stage of hardening in outdoor-grown seedlings. Frost hardiness developed at a constant rate in response to a shortening photoperiod once it had declined to about 12 h. This rate was consistent with the hardening rate that occurred in outdoor-grown seedlings. Hardening stopped when the photoperiod became constant, indicating a tight coupling between changes in photoperiod and hardiness development. When the photoperiod was held constant, the extent of frost hardiness was directly dependent on the photoperiod but the rate of hardening was apparently independent of the length of photoperiod. However, the rate of hardening was dependent on the rate at which the photoperiod shortened, increasing linearly with increases in the rate of change in photoperiod between 0 and 3 min d^?1. These results suggest shortening photoperiods control the first stage of the hardening process by regulating the rate of hardening. Frost hardening was inherently unstable once the maximum hardiness was reached since spontaneous dehardening occurred in spite of the controlled conditions. Dehardening also occurred when the photoperiod was lengthened suggesting that the cue for dehardening to commence was the shift from shortening to lengthening photoperiods.     

Acute toxicity of Cd in stream invertebrates in relation to pH and test design

The acute toxicity of Cd to three stream invertebrates (Baetis rhodani Pict., Leptophlebia marginata (L.) and Pisidium sp.) was tested at pH 5 and 7 simultaneously in static (ST) and flow through (FT) systems. In the static design, the animals were kept individually in small boxes containing aerated stream water. In the flow through system, the three species were kept together in circular aquaria simulating stream ecosystems with patches of gravel and recirculating aerated stream water. The nominal Cd concentrations ranged from 0 to 5 mg 1^–1 during an exposure period of 120 h.The graphically obtained LC₅₀ (120 h) values for B. rhodani were: pH 7: 2.3 mg 1^–1 (ST), 2.5 mg 1^–1 (FT) and pH 5: 3 mg 1^–1 (ST), 1 mg 1^–1 (FT). For L. marginata the following values were found: pH 7: > 5 mg 1^–1 (ST), 4.4 mg 1^–1 (FT) and pH 5: > 5 mg 1^–1 (ST), 3.6 mg 1^–1 (FT). Pisidium sp. showed a 100% survival in all Cd- and pH-treatments. From thee values it can be concluded that L. marginata is more tolerant than B. rhodani and that especially both species tolerated Cd better in the ST system than in the FT system, especially at pH 5.At neutral pH the Cd-concentrations in the animals were higher than at low pH, probably due to surface adsorption of Cd onto the body. B. rhodani tended to molt more at low than at neutral pH.     

Coastal sand dune stabilization in the Pacific Northwest

Coastal dunes are stabilized in three stages: (1) The initial stage uses sand-stilling grasses established vegetatively. For this purpose, European beachgrass,Ammophila arenaria (L.) Link., is most used, followed by American beachgrass,A. breviligulata Fern., or American dunegrass,Elymus mollis Trin. Large solid plantings must be made with the spacing and number of plants per hill adjusted to the site conditions. Plantings, using Nitrogen fertilizer, are made during the dormant season. (2) Secondary permanent grasses and legumes are seeded 2 years later when sand movement slows and the initial grasses begin to lose their vigor. Most used are seaside lupine,Lupinus littoralis Dougl., purple beachpea,Lathyrus japonicus Willd., seashore bluegrass,Poa macrantha Vasey., and native red fescue,Festuca rubra L. (3) Final control is with woody plants that are well adapted to rough areas. Scotch broom,Cytisus scoparius (L.) Link. Hooker willow,Salix hookeriana Barratt, Nootka rose,Rosa nutkana Prese., and shore pine Pinus contorta Dougl., have been extensively used.Presented at the Seventh International Biometeorological Congress, 17–23 August 1975, College Park, Maryland, U.S.A.     

Mycorrhizal status of herb-layer plants in a fertilized oak-pine forest

In the years 1986–1988 investigations were carried out on the influence of fertilizers on the mycorrhizal status of herb-layer plants of a mixed oak-pine forest in the Niepolomice Forest (Southern Poland). The site enrichment with N, P, K altered the frequency of mycorrhizal infection in most species investigated and decreased the percentage of colonized root cells. Some species disappear after fertilization (e.g. Vaccinium myrtillus L.). Among the species vigorously expanding after the treatment, mainly nonmycorrhizal plants (e.g. Festuca gigantea (L.) Vill.) were found. Some species (e.g. Milium effusum L.) favourably reacting to mineral fertilization, lost their mycorrhizae.     

The forth estuary: a nursery and overwintering area for North Sea fishes

The fish community of the Forth estuary, Scotland, has several components — estuarine resident species, diadromus migratory species, marine and freshwater adventitious species, marine juvenile migrants using the area as a nursery, and adults of marine species with seasonal migrations. The population changes during the period 1981–1988 in six species representative of the last two categories are described here. The species are the juvenile gadoids, whiting, Merlangius merlangus (L.) and cod, Gadus morhua L., juvenile flatfish, plaice, Pleuronectes platessa L. and common dab, Limanda limanda (L.), and the adult clupeids, sprat, Sprattus sprattus (L.) and herring, Clupea harengus L.The species' population changes are described in relation to aspects of wider significance such as commercial fishing for these species in North Sea areas. In particular, the underlying decline of the juvenile cod populations since 1981 in the Forth Estuary is discussed in relation to the recent reduction in commercial fishing quotas.     

The efficiency of evapotranspiration of landfill leachate in the soil–plant system with willow Salix amygdalina L.

The application of a specific species of willow—Salix amygdalina L., marked by high transpiration ability—is a cheap and effective method of landfill leachate disposal. A 2-year study examined the effectiveness of leachate evapotranspiration from soil–plant systems with willow species S. amygdalina L. Evapotranspiration from soil–plant systems planted with willow was from 1.28 up to 5.12 times higher than evaporation from soil surface barren of vegetation. This proves the usefulness of soil–plant systems with willow in landfill leachate treatment through vaporization. Evapotranspiration efficiency, as opposed to total amount of water added into the lysimeter, was not strong enough to vaporize all input of the landfill leachate in the lysimeters. This may indicate that the ground water requires isolation when soil systems remain under landfill leachate irrigation. Linear dependence between willow biomass growth and transpiration was observed to be significant (p < 0.05). Additionally, the research showed that the application of sewage sludge into the soil caused an increase in vaporization efficiency.     

Post-fire resprouting of shrubby species in Mediterranean maquis

The vegetative resprouting of mediterranean maquis shrubby species was examined eight years after fire. Post-fire regeneration occurs through the resprouting of stumps. All species (Quercus ilex L., Phillyrea latifolia L., Arbutus unedo L., Erica arborea L., Pistacia lentiscus L.) show a rapid growth in the first years after fire, and a decrease already from 4^th–5^th year.All the species survived the fire and reconstituted a community similar to that unburned in a relatively short time span.     

Root Strength and Root Area Ratio of Forest Species in Lombardy (Northern Italy)

Forest vegetation is known to increase hillslope stability by reinforcing soil shear resistance and by influencing hydrologic conditions of soil. Although the importance of plant root systems for hillslope stability has received considerable attention in recent years, the quantification of such an effect needs more investigation. In this paper, we present a synthesis of the data gathered in the last 5 years for some species in different locations of the Alps and Prealps of Lombardy (Northern Italy) with the aim to increase our knowledge on root tensile strength and on Root Area Ratio distribution within the soil. Concerning root tensile strength we developed tensile strength–diameter relationships for eight species: green alder (Alnus viridis(Chaix) D.C.), beech (Fagus sylvatica L.), red willow (Salix purpurea L.), goat willow (Salix caprea L.), hazel (Corylus avellana L.), European ash (Fraxinus excelsior L.), Norway spruce (Picea abies (L.) Karst.) and European larch (Larix decidua Mill.). Results show a great variability among the different species and also for the same species. In general, however, root strength (in terms of tension) tends to decrease with diameter according to a power law, as observed by other Authors. Comparing the power law fitting curves for the considered species, it can be observed that they fall in a relatively narrow band, with the exception of hazel, which appears the most resistant. Concerning the evaluation of root distribution within the soil we estimated the Root Area Ratio (the ratio between the area occupied by roots in a unit area of soil) according to its depth for five species (beech, Norway spruce, European larch, mixed hazel and ash) in three locations of Lombardy. Results show that there is a great variability of root density for the same species well as for different points at the same locality. The general behaviour of root density, in any case, is to decrease with depth according to a gamma function for all the studied species. The results presented in this paper contribute to expanding the knowledge on root resistance behaviour and on root density distribution within the soil. The studied location have allowed the implementation of soil–root reinforcement models and the evaluation of the vegetation contribution to soil stability.     

Differences in uptake kinetics of ammonium and nitrate in legumes and cereals

Influx isotherms were obtained for nitrate and ammonium from three legumes, Cajanus cajan (L.) Millsp., Cicer arietinum L. and Arachis hypogaea L. and three cereals, Sorghum bicolor (L.) Moench., Pennisetum glaucum L. and Zea mays L. The transition in influx isotherms for both nitrogen sources was found to be within the concentration range (0.05–2.5 mM) tested. There were significant differences in Km and Vmax for ammonium between legumes and cereals. The difference in the kinetic properties for nitrate uptake between the two groups of plants only became apparent at the higher concentration tested. Legumes translocated absorbed nitrate and ammonium to shoots more rapidly than cereals. Results show that there are significant differences in uptake and translocation of ammonium and nitrate between legumes and cereals.     

Effects of climatic warming on cold hardiness of some northern woody plants assessed from simulation experiments

Effects of climatic warming on cold hardiness were investigated for some northern woody plants. In the first experiment, seedlings of Norway spruce ( Picea abies [L.] Karst.), Scots pine ( Pinus sylvestris L.) and lodgepole pine ( Pinus contorta Dougl. var. latifolia Engelm.) were exposed to naturally fluctuating temperatures averaging −6°C (ambient) and 0°C (elevated) for 16 weeks in midwinter before they were thawed and re-saturated with water. In lodgepole pine, needle sugar concentrations had decreased by 15%, and the temperature needed to induce 10% injury to needles in terms of electrolyte leakage had increased by 6°C following treatment to elevated as compared with control temperatures. In contrast, Norway spruce and Scots pine showed no effects. The lack of an effect for Scots pine was ascribed to seedlings containing unusually large energy reserves that buffered respiratory expenditure of sugars. A strong, linear relationship between levels of cold hardiness, assessed by the electrolyte leakage method, and sugars was found when combining data from this and previous, similar experiments. In the second experiment, the evergreen dwarf shrub Empetrum hermaphroditum Hagerup was analysed for leaf cold hardiness, using the electrolyte leakage method, and sugar concentrations in late spring and late autumn during the third year of a warming experiment in a subarctic dwarf shrub community. The objective was to test the hypothesis that warming in the growing season alters hardening/dehardening cycles by increasing soil nitrogen mineralization and plant growth. Data found, however, suggested that cold hardening/dehardening cycles were unaffected by warming.     

Elevated CO2 and the mineral content of herbaceous and woody plants

The CO₂ enrichment effects (300–650 µmol mol^-1) on mineral concentration (N, P, K, Ca, Mg, Mn, Fe, Zn), absolute total mineral contents per individual and of whole stands of four herbaceous (Trifolium repens L.,Trifolium pratense L.,Lolium perenne L.,Festuca pratensis HUDS.) and two woody species (Acer pseudo-platanus L.,Fagus sylvatica L.) were investigated.In general, the mineral concentration of the plant tissues decreased (all six species: N>Ca>K>Mg) with the exception of P. Mn and Fe were only determined for the tree species. Both decreased in concentration (Mn>Fe). Zn was only analysed forTrifolium pratense andFestuca pratensis and decreased significantly in the grass.Despite of decreases in concentrations of as much as 20% in some cases there were increases in absolute amounts per individual and, therefore, in the whole vegetation up to 25% because of the enhanced dry matter accumulation at elevated CO₂ supply.Dedicated to Prof. Dr. R. Bornkamm, TU-Berlin, on behalf of his 60th birthday     

Soil seed banks of woodland, heathland, grassland, mire and montane communities, Cairngorm Mountains, Scotland

The size and species composition of soil seed banks were assessed at 111 altitudinally diverse sites in the Cairngorm Mountains. Mean densities of germinable seeds varied from 83 000 m^–2 in Scots pine (Pinus sylvestris L.) woodland at 230–490 m to 200 m^–2 in moss (Racomitrium lanuginosum (Hedw.) Brid.) heath at 1000–1120 m. Seed banks were dominated by Calluna vulgaris (L.) Hull, not only wherever it was prominent in the vegetation, but also at some sites with less than 5% cover of parent plants in the ground vegetation. Many species conspicuous in the vegetation were under-represented in or absent from the seed bank and surface vegetation generally was more species rich than was the underlying seed bank, especially in high montane communities. Multiple regression was used to examine the relationship between the density of buried Calluna seeds and the abundance of parent plants in the vegetation, site altitude and the organic matter content of the soil. The model fitted to woodland communities accounted for 95% of the variation in seed density. The heathland model was less predictive but still explained 52% of the variation in seed bank size. In mire communities there was no relationship, collective or individual, between buried seed density and the measured environmental variables, possibly due to variations in the duration and frequency of waterlogging at these sites. The potential role of seed banks for initiating the recolonisation of disturbed ground is discussed. Densities of buried seeds at most Calluna-dominant sites were probably sufficient to generate successful recolonisation but the prospects for recovery were poor at other sites, particularly in graminaceous communities at 800 m or higher.