Determinants of woody species richness in Scot pine and beech forests: climate, forest patch size and forest structure

We analysed patterns of woody species richness in Pinus sylvestris and Fagus sylvatica forests in Catalonia (NE Spain) from forestry inventory databank in relation to climate and landscape structure. Both types of forests are found within the same climatic range, although they have been managed following somewhat different goals. Overall, woody species richness significantly increased when conditions get closer to the Mediterranean ones, with milder temperatures. Differences between the two types of forests arose when comparing the relationship between richness and forest patch size. Woody species richness increased in pine forests with patch size, while the opposite trend was observed in beech forests. This pattern is explained by the different behaviour of structural canopy properties, since leaf area index and canopy cover showed a steeper increase with increasing forest patch size in Fagus forests than in Pinus ones. Accordingly, richness decreased with canopy cover in Fagus plots, but not in Pinus ones. We suggest that these differences would be related to management history, which may have enhanced the preservation of beech stands in larger forest landscape units.     

Nitrogen metabolism of Douglas fir and Scots pine as affected by optimal nutrition and water supply under conditions of relatively high atmospheric nitrogen deposition

Nitrogen metabolism of the needles of 40-year-old Douglas fir and Scots pine trees, growing in two forest stands on cation-poor and acidic sandy soil with a relatively high atmospheric nitrogen deposition was studied. The composition of the free amino acid (FAA) pool, the concentrations of total nitrogen and soluble protein and the activities of glutamine synthetase (GS) and glutamate dehydrogenase (GDH) were determined in the needles. An excessive nitrogen supply by a high atmospheric nitrogen deposition in both forest stands was indicated by the high concentrations of total nitrogen and the amino acids arginine, glutamic acid, glutamine and aspartic acid in control trees. In addition the effect of optimal nutrition and water supply (fertigation) on the needle nitrogen metabolism was evaluated. The total concentration of the FAA pool in needles of both tree species was lower in the fertigated than in the non-fertigated (control) trees, except for 1-year-old needles of Scots pine, in which the concentration after fertigation did not differ from the control. The lower total FAA concentration in the fertigated trees could be attributed to arginine, the concentration of which was on average 60% lower than in the control. Neither the concentration of soluble protein nor the activity of GS were influenced by fertigation. The activity of GDH in fertigated trees only differed significantly from the control in October. Scots pine needles had higher concentrations of protein (50%) and higher activities of GS (44%) and GDH (25%) than Douglas fir needles. Possible explanations for the lower vitality of Douglas fir compared to Scots pine are given.     

The retention of organic matter in soils

The turnover of C in soils is controlled mainly by water regimes and temperature, but is modified by factors such as size and physicochemical properties of C additions in litter or root systems, distribution of C throughout the soil as root systems, or addition as litter, distribution of C within the soil matrix and its interaction with clay surfaces.Soil factors which retard mineralization of C in soils are identified from correlations of C contents of soils with other properties such as clay content and base status. The rate and extent of C mineralization depends on the chemistry of the added organic matter and interaction with clays of the microbial biomass and metabolites.The organomineral interactions are shown to depend on cation bridges involving mainly Ca in neutral to alkaline soils, Al in acid soils and adsorption of organic materials on iron oxide surfaces. The various organomineral interactions lead to aggregations of clay particles and organic materials, which stabilizes both soil structure and the carbon compounds within the aggregates.     

Above- and belowground organic matter storage and production in a tropical pine plantation and a paired broadleaf secondary forest

The distribution of tree biomass and the allocation of organic matter production were measured in an 11-yr-old Pinus caribaea plantation and a paired broadleaf secondary forest growing under the same climatic conditions. The pine plantation had significantly more mass aboveground than the secondary forest (94.9 vs 35.6 t ha^-1 for biomass and 10.5 vs 5.0 t ha^{-1 for litter}), whereas the secondary forest had significantly more fine roots (⩽2 mm diameter) than the pine plantation (10.5 and 1.0 t ha^-1, respectively). Standing stock of dead fine roots was higher than aboveground litter in the secondary forest. In contrast, aboveground litter in pine was more than ten times higher than the dead root fraction. Both pine and secondary forests had similar total organic matter productions (19.2 and 19.4 t ha^-1 yr^-1, respectively) but structural allocation of that production was significantly different between the two forests; 44% of total production was allocated belowground in the secondary forest, whereas 94% was allocated aboveground in pine. The growth strategies represented by fast growth and large structural allocation aboveground, as for pine, and almost half the production allocated belowground, as for the secondary forest, illustrate equally successful, but contrasting growth strategies under the same climate, regardless of soil characteristics. The patterns of accumulation of organic matter in the soil profile indicated contrasting nutrient immobilization and mineralization sites and sources for soil organic matter formation.     

Controls on the dynamics of dissolved organic carbon and nitrogen in a Central European deciduous forest

Despite growing attention concerning therole of dissolved organic matter (DOM) inelement cycling of forest ecosystems, thecontrols of concentrations and fluxes of bothdissolved organic carbon (DOC) and nitrogen(DON) under field conditions in forest soilsremain only poorly understood. The goal ofthis project is to measure the concentrations and fluxes of DON, NH₄ ⁺, NO₃ ^–and DOC in bulkprecipitation, throughfall, forest floorleachates and soil solutions of a deciduousstand in the Steigerwald region (northernBavaria, Germany). The DOC and DONconcentrations and fluxes were highest inleachates originating from the Oa layer of theforest floor (73 mg C L^–1, 2.3 mg NL^–1 and about 200–350 kg C, 8–10 kg Nha^–1 yr^–1). They were observed to behighly variable over time and decreased in themineral topsoil (17 mg C L^–1, 0.6 mg NL^–1 and about 50–90 kg C, 2.0 to 2.4 kg Nha^–1 yr^–1). The annual variability ofDOC and DON concentrations and subsequentialDOC/DON ratios was substantial in allsolutions. The DOC and DON concentrations inthroughfall were positively correlated withtemperature. The DOC and DON concentrationsdid not show seasonality in the forest floorand mineral soil. Concentrations were notrelated to litterfall dynamics but didcorrespond in part to the input of DOC and DONfrom throughfall. The throughfall contributionto the overall element fluxes was higher forDON than for DOC. Concentrations and fluxes ofDON were significantly correlated to DOC inthroughfall and the Oi layer. However, thecorrelation was weak in Oa leachates. Inaddition, seasonal and annual variation ofDOC/DON ratios indicated different mechanismsand release rates from the forest floor forboth components. The concentrations of DOC andDON in forest floor leachates were in mostcases dependent neither on the pH value orionic strength of the solution, nor on thewater flux or temperature changes. As aconsequence, the DOC and DON fluxes from theforest floor into the mineral soil werelargely dependent on the water flux if annualand biweekly time scales are considered.     

Depth distribution and composition of seed banks under different tree layers in a managed temperate forest ecosystem

In the present work we examined the composition and distribution across three soil layers of the buried soil seed bank under three different overstory types (Fagus sylvatica, Quercus robur, Pinus sylvestris) and in logging areas in a 4383-ha forest in central Belgium. The objectives were: (1) to investigate whether species composition and species richness of soil seed banks are affected by different forest stands; (2) to examine how abundant are habitat-specific forest species in seed banks under different planted tree layers. The study was carried out in stands which are replicated, managed in the same way (even-aged high forest), and growing on the same soil type with the same land-use history. In the investigated area, the seed bank did show significant differences under oak, beech, pine and in logging areas, respectively in terms of size, composition and depth occurrence. All species and layers taken together, the seed bank size ranked as follows: oakwood > beechwood > logging area > pinewood. The same pattern was found for forest species. Seed numbers of Betula pendula, Calluna vulgaris, Dryopteris dilatata and Rubus fruticosus were significantly higher under the beech canopy. Carex remota, Impatiens parviflora and Lotus sp. showed a significantly denser seed bank in logging areas, while Digitalis purpurea seeds were significantly more abundant in soils under the oak canopy. The fact that the seed bank of an originally homogeneous forest varies under different planted stands highlights that a long period of canopy conversion can affect the composition and depth of buried seeds.     

森林类型对土壤有机质、微生物生物量及酶活性的影响

以澳大利亚南昆士兰州典型森林类型——湿地松、南洋杉和贝壳杉林为对象,开展土壤可溶性有机碳和氮(SOC和SON)、微生物生物量碳和氮(MBC和MBN),以及土壤酶活性的研究,剖析森林类型对土壤质量的影响.结果表明:不同林型土壤SOC、SON含量分别在552 ～1154 mg·kg-1和20.11～57.32mg·kg-1;MBC、MBN分别在42～149 mg·kg-1和7～35 mg·kg-1.MBC、MBN之间呈显著相关.土壤几丁质酶、酸性磷酸酶、碱性磷酸酶和β-葡萄糖苷酶的活性分别为2.96 ～7.63、16.5 ～29.6、0.79 ～ 3.42和3.71 ～9.93 μg ·g-1·h-1,亮氨酸氨肽酶活性为0.18～0.46 μg·g-1·d-1.不同林型土壤SOC含量,以及土壤几丁质酶和亮氨酸氨肽酶活性为湿地松林、南洋杉林、贝壳杉林依次降低;而SON含量为南洋杉林＞贝壳杉林＞湿地松林,且南洋杉林的SON含量显著(P＜0.05)高于湿地松林;MBC和MBN以及碱性磷酸酶活性为贝壳杉林＞湿地松林＞南洋杉林;酸性磷酸酶和β-葡萄糖苷酶活性为湿地松林＞贝壳杉林＞南洋杉林.在土壤生物代谢因子中,MBC、MBN、SON和亮氨酸氨肽酶对不同森林类型土壤影响较大.     

Decomposition of organic matter from 36 soils in a long-term pot experiment

The organic matter contents of thirty-six soils were measured annually for twenty years in a pot experiment. The soils originated mainly from arable land and varied in initial organic matter content, texture and pH. The soils were stored at an average air temperature of around 13 °C and every year each soil was mixed thoroughly. Throughout the experiment, soil moisture was kept between 50-70% of its water holding capacity. No organic matter was added during the experiment, so that gross soil organic matter decomposition could be assessed. Relative decomposition rates of soil organic matter decreased as time proceeded. Despite the wide range of soils studied, it was found that during the initial decades, the pattern of soil organic matter degradation was strongly correlated with the organic matter content of the soils at the start of the experiment. This means that during this period the time course of the organic matter content of the soils in our experiment can be estimated from the initial soil organic matter content alone.     

Dynamics of organic matter in soils

Summary Dynamics of C, N, S, and to some extent P are expressed by a knowledge of the size and turnover rates of plant constituents such as soluble C and N components, cellulose and hemicellulose, and lignin. Soil organic matter constituents include: the microbial biomass as determined chemically or microscopically, non-biomass active components determined by isotopic dilution, stabilized N constituents for which good techniques are not yet available, and resistant or old C and associated N determined by carbon dating. The processes involved in the nutrient transformations and transfers are reasonably well understood. The control mechanisms require further elucidation to be able to extrapolate from the laboratory to the field, and between field sites. Major control mechanisms requiring further insight include the effects of C availability on transformations of C and N. The other control for which every little is known is that of spatial compartmentalization. Compartmentalization ranges from landscape or management sequences to pedogenic layers, rhizosphere-mycorrhizal effects, clay-sesquioxide surfaces, aggregation, localized enzymes, and microbial effects such as membrane boundaries. Control mechanisms for concurrent mineralization-immobilization, the stabilization of microbial products, and the relative role of the biomass as a catalyst rather than as a source-sink for nutrients, must be understood. There is potential for combining a knowledge of microbial production and turnover with that of the roles of the soil organic active fraction as a temporary storehouse for nutrients. This, in conjunction with management techniques such as zero tillage and crop rotation, should make it possible to better utilize soil and fertilizer N, especially in areas of the world where the cost of nutrients is high relative to the value of the crop grown.Introductory lecture     

Sorption of dissolved organic matter by mineral soils of the Siberian forest tundra

Because of low net production in arctic and subarctic surface water, dissolved organic matter (DOM) discharged from terrestrial settings plays an important role for carbon and nitrogen dynamics in arctic aquatic systems. Sorption, typically controlling the export of DOM from soil, may be influenced by the permafrost regime. To confirm the potential sorptive control on the release of DOM from permafrost soils in central northern Siberia, we examined the sorption of DOM by mineral soils of Gelisols and Inceptisols with varying depth of the active layer. Water‐soluble organic matter in the O horizons of the Gelisols was less (338 and 407 mg C kg^?1) and comprised more dissolved organic carbon (DOC) in the hydrophobic fraction (HoDOC) (63% and 70%) than in the O horizons of the Inceptisols (686 and 706 mg C kg^?1, 45% and 48% HoDOC). All A and B horizons from Gelisols sorbed DOC strongly, with a preference for HoDOC. Almost all horizons of the Inceptisols showed a weaker sorption of DOC than those of the Gelisols. The C horizons of the Inceptisols, having a weak overall DOC sorption, sorbed C in the hydrophilic fraction (HiDOC) stronger than HoDOC. The reason for the poor overall sorption and also the preferential sorption of HiDOC is likely the high pH (pH>7.0) of the C horizons and the smaller concentrations of iron oxides. For all soils, the sorption of HoDOC related positively to oxalate‐ and dithionite–citrate‐extractable iron. The A horizons released large amounts of DOC with 46–80% of HiDOC. The released DOC was significantly (r=0.78, P<0.05) correlated with the contents of soil organic carbon. From these results, we assume that large concentrations of DOM comprising large shares of HiDOC can pass mineral soils where the active layer is thin (i.e. in Gelisols), and enter streams. Soils with deep active layer (i.e. Inceptisols), may release little DOM because of more frequent infiltration of DOM into their thick mineral horizons despite their smaller contents of reactive, poorly crystalline minerals. The results obtained for the Inceptisols are in agreement with the situation observed for streams connecting to Yenisei at lower latitudes than 65°50′ with continuous to discontinuous permafrost. The smaller sorption of DOM by the Gelisols is in agreement with the larger DOM concentrations in more northern catchments. However, the Gelisols preferentially retained the HoDOC which dominates the DOC in streams towards north. This discrepancy can be explained by additional seepage water from the organic horizons that is discharged into streams without intensive contact with the mineral soil.     

Soil organic matter dynamics under decaying wood in a subtropical wet forest: effect of tree species and decay stage

Decaying wood is an important structural and functional component of forests: it contributes to generate habitat diversity, acts as either sink or source of nutrients, and plays a preponderant role in soil formation. Thus, decaying wood might likely have measurable effects on chemical properties of the underlying soil. We hypothesized that decaying wood would have a stronger effect on soil as decomposition advances and that such effect would vary according to wood quality. Twenty logs from two species with contrasting wood properties (Dacryodes excelsa Vahl. and Swietenia macrophylla King) and at two different decay stages (6 and 15 years after falling) were selected, and soil under and 50 cm away from decaying logs was sampled for soil organic matter (SOM) fractions [NaOH-extractable and water-extractable organic matter -(WEOM)] and properties (WEOM aromaticity). NaOH-extractable C and WEOM were higher in the soil influenced by 15-year-old logs, while the degree of aromaticity of WEOM was higher in the soil influenced by the 6-year-old logs. Decaying logs did influence properties of the underlying soil with differing effects according to the species since there was more NaOH-extractable C in the soil associated to D. excelsa logs and more WEOM in the soil associated to S. macrophylla older logs. It is proposed that such effects occurred through changes in the relative quantity and quality of different SOM fractions, as influenced by species and advancement in decomposition. Through its effect on SOM and nutrient dynamics, decaying wood can contribute to the spatial heterogeneity of soil properties, and can affect process of soil formation and nutrient cycling. Responsible Editor: Barbara Wick.     

土地利用方式对土壤有机质的影响

通过对地带性常绿阔叶林、杉木人工林、农田、竹林等不同土地利用方式下土壤有机质总量、活性有机质及其组分的研究,发现土地利用方式对土壤有机质和活性有机质各组分的影响差异显著,其中阔叶林含量最高,杉木人工林低于阔叶林,竹林和农田最低。这些差别主要是由于凋落物的数量、质量以及各种管理措施不同所致。     

Response of Douglas-fir and amabilis fir to split-root nutrition with inorganic and organic nitrogen

There is limited understanding of the spatial plasticity of conifer root growth in response to inorganic and organic nitrogen (N). In this study, slow-growing amabilis fir and fast-growing Douglas-fir, and slow- and fast-growing seedlots of the latter species were examined for their ability to proliferate roots preferentially in compartments of sand/peat medium enriched in organic and inorganic forms of N. In one experiment, N was supplied as 7.1 or 0.71 mM ammonium, nitrate and ammonium nitrate, and in a second experiment, N was supplied as ammonium or glycine. The seedlings’ ability to compensate for the starvation of a portion of the root system was assessed by measuring biomass of leaves, stems and roots, and foliar N concentration. Both fast- and slow-growing seedlots of Douglas-fir and slow-growing amabilis fir were able to proliferate roots in compartments of soil enriched with inorganic and organic N. In the first experiment, whole plant and root biomass was greatest when N was provided as ammonium followed by nitrate, and in the second experiment, seedling whole and root biomasses did not differ between ammonium and glycine treatments. All seedlings were able to compensate for the starvation of a portion of the root system, thus total plant biomass did not differ between split-root treatments; however, foliar N contents were lower in the 7.1/0.71 mM inorganic N split-root treatments. Foliar N concentrations were also lower in seedlings supplied with glycine.     

Influence of root temperature on growth of Pinus sylvestris,Fagus sylvatica,Tilia cordata and Quercus robur

One-year-old tree seedlings were incubated in a greenhouse from April to July, under natural daylight conditions, with their root systems at constant temperatures of 5, 10, 15, 20, 25, 30 and 35 °C and with the above ground parts kept at a constant air temperature of 18–20 °C. The course of height growth, total mass increment, root, shoot and leaf weight as well as leaf areas were measured. The results indicate that clear differences exist in the optimal root zone temperatures for various growth parameters in different tree species. Pinus sylvestris had a maximal height increment at about 5–10 °C and maximal total mass increment at 15 °C root temperature. In contrast, the optimum for Quercus robur was at 25 °C. Tilia cordata and Fagus sylvatica had their optima for most growth parameters at 20 °C. The root temperature apparently indirectly influenced photosynthesis (dry weight accumulation) and respiration loss. From the observed symptoms and indications in the literature it seems probable that a change in hormone levels is involved as the main factor in the described effects. Variation of root temperature had only an insignificant effect on bud burst and the time at which the shoots sprouted. Apparently species of northern origin seem to have lower root temperature optima than those of more southern origin. This is to be verified by investigation of other tree species.     

Anaerobic induced ethanol synthesis in the stems of greenhouse-grown conifer seedlings

Ethanol synthesis was induced in stem segments from greenhouse-grown conifer seedlings by placing them in a N₂ atmosphere at 30 °C for 24 h. Stems from ponderosa pine,Pinus ponderosa Dougl. ex Laws., sugar pine,Pinus lambertiana Dougl., Pacific silver fir,Abies amabalis Dougl. ex Forbes, and lodgepole pine,Pinus contorta Dougl. ex Loud, produced the highest quantities of ethanol. This group also had the smallest and slowest growing stems. Within each of these species the amount of ethanol produced was inversely related to the stem volume. Stems from western hemlock,Tsuga heterophylla (Raf.) Sarg., grand fir, Abies grandis Dougl. ex Forbes, Douglas-fir,Pseudotsuga menziesii (Mirb.) Franco, and western redcedar,Thuja plicata Donn ex D. Don, all produced equivalent but low ethanol concentrations. These species had the largest and fastest growing stems. In this group only grand fir exhibited an inverse relationship between ethanol concentrations and stem volume. The relative amounts of ethanol synthesized by stems from Douglas-fir, western hemlock and western redcedar seedlings were not the same as subsequently observed in logs from mature trees of the same species under field conditions. Differences in the anaerobic environments for the two stem types could have affected the quantities of ethanol produced. The observed high amounts of ethanol produced by the stems from pine species were discussed in terms of their ability to handle periods of anaerobic stress or hypoxia.     

Copper uptake by some forest tree species from an acid sandy soil

Summary The behaviour of some young conifers and broadleaves on a very acid sandy humuspodzol soil was studied in pot trials. Species differed considerably in foliar copper concentration, reaction to N fertilization and development of Cu deficiency symptoms. Foliar copper concentrations were highest for the broadleaves (esp. oak and beech), and was increased by Cu and N fertilization, the last reaction showing an optimum curve. Cu deficiency symptoms could easily be induced in the conifers but not in the broadleaves.     

Root biomass and rooting profile of naturally regenerated beech in mid-elevation Scots pine woodlands

The natural establishment of shade-tolerant forest species such as beech (Fagus sylvatica L.) occurs in naturally regenerated Scots pine (Pinus sylvestris L.) woodlands that develop on former pastures and cultivated lands. To examine possible effects of underground competition in beech establishment, we studied the root biomass and the rooting profile of 53 mixed Scots pine-natural beech woodlands in French mid-elevation volcanic areas. Stands were arranged along a maturation gradient. Roots were sampled using the root-auger technique (0 to 75 cm depth every 15 cm). In addition, 23 young beeches were uprooted to study the entire root system. Total beech fine-root biomass was closely correlated with most beech aerial characteristics, (e.g., height, diameter and girth), and correlated moderately with tree age. However, it correlated poorly with basic competition indices such as stand density and basal area. Conversely, competition indices including vertical dimensions of competing trees were correlated with the underground biomass, probably as a result of redundancy with beech height. The rooting profile (fine roots, < 5 mm) of beech and pine were quite similar, and did not change significantly along the stand maturation gradient. Beech has a heart-shaped root system while pine is more plate-like and dimorphic. Beech fine-root biomass progressively surpassed pine biomass throughout the soil layers, thus confirming that it is dynamic and competitive in mature mixed stands. The coexistence of the root systems of beech and pine in the same soil layers presumably results in strong underground competition.     

The occurrence of flowering and fruiting on individual trees over 3 years and their effects on subsequent crown condition

Summary The level of fruiting in four forest trees species (Picea sitchensis, P. abies, Pinus sylvestris and Fagus sylvatica) was monitored in Great Britain over the period 1989–1991. In addition, assessments of crown transparency were available for many of the trees for 1987 and 1988. The monitoring period encompassed severe summer droughts in 1989 and 1990, with wetter conditions in 1991. Variations in the level of fruiting in spruce and beech (Fagus sylvatica L.) were seen, with a marked peak in 1990. No pattern was apparent in Scots pine (Pinus sylvestris L.). Coning, which was greater in trees with the least transparent crowns, had no discernible effect on the crown transparency of the conifers. Cupule production in beech was greatest in trees with the most transparent crowns, and trees with high numbers of cupules in 1990 tended to have greater crown dieback recorded in 1991.     

Soil organic matter and litter chemistry response to experimental N deposition in northern temperate deciduous forest ecosystems

The effects of atmospheric nitrogen (N) deposition on organic matter decomposition vary with the biochemical characteristics of plant litter. At the ecosystem‐scale, net effects are difficult to predict because various soil organic matter (SOM) fractions may respond differentially. We investigated the relationship between SOM chemistry and microbial activity in three northern deciduous forest ecosystems that have been subjected to experimental N addition for 2 years. Extractable dissolved organic carbon (DOC), DOC aromaticity, C : N ratio, and functional group distribution, measured by Fourier transform infrared spectra (FTIR), were analyzed for litter and SOM. The largest biochemical changes were found in the sugar maple–basswood (SMBW) and black oak–white oak (BOWO) ecosystems. SMBW litter from the N addition treatment had less aromaticity, higher C : N ratios, and lower saturated carbon, lower carbonyl carbon, and higher carboxylates than controls; BOWO litter showed opposite trends, except for carbonyl and carboxylate contents. Litter from the sugar maple–red oak (SMRO) ecosystem had a lower C : N ratio, but no change in DOC aromaticity. For SOM, the C : N ratio increased with N addition in SMBW and SMRO ecosystems, but decreased in BOWO; N addition did not affect the aromaticity of DOC extracted from mineral soil. All ecosystems showed increases in extractable DOC from both litter and soil in response to N treatment. The biochemical changes are consistent with the divergent microbial responses observed in these systems. Extracellular oxidative enzyme activity has declined in the BOWO and SMRO ecosystems while activity in the SMBW ecosystem, particularly in the litter horizon, has increased. In all systems, enzyme activities associated with the hydrolysis and oxidation of polysaccharides have increased. At the ecosystem scale, the biochemical characteristics of the dominant litter appear to modulate the effects of N deposition on organic matter dynamics.