Enchytraeid numbers,biomass and respiratory metabolism in a beech woodland - Wytham Woods,Oxford

Summary The mean annual population density of enchytraeids in the litter and upper 6 cm of soil was found to be 14,590 m^-2. Mean individual weights approximated 116, 158 and 151 g wet wt. in the litter, 0–3 cm and 3–6 cm strata respectively. The mean biomass was estimated to be 1.908 g wet wt m^-2. Vertical migration was shown to occur, the upward migration in late Autumn was a response to temperatures in the litter being temporarily higher than those of the 0–3 cm stratum. As a result of upward vertical migration and possibly recruitment, numbers reached a maximum in the litter during Winter. On an annual basis the litter, 0–3 and 3–6 cm strata contained 41.43, 46.44 and 12.12% respectively of the extracted enchytraeids. The equivalent biomass values were 33.18, 52.46 and 14.36%. Total numbers and biomass revealed a general picture of high values in late Autumn to Winter which gradually decreased through Spring and Summer except for a minor peak in May–June. The annual oxygen consumption of the enchytraeids approximated 4.285 l O₂ m^-2 yr^-1 (20.461 kcal85.610 kJ) and accounted for 1.63% of the total soil metabolism. A production/biomass (P/B) ratio of 4.93 was estimated as was a net population efficiency of 32%. For this site the contribution by enchytraeids to total soil respiration is about one-third that of the earthworms and an order of magnitude greater than that of the nematodes.     

The fleas of a population of weasels in Wytham Woods, Oxford

Weasels were regularly livetrapped, anaesthetized and searched for fleas over two years, in Wytham Woods, near Oxford. Eleven species of fleas were identified among 262 collected. Seven normally occur on small rodents, one on rats, two on moles and one on birds. Fleas were collected on 103 of 338 occasions when an unconscious weasel was examined; on average males carried 0·6 fleas, females 1·5 fleas, but there was no significant difference between the proportion of males and females infested. The species distribution of the fleas on weasels in Wytham suggests that they acquire their fleas mainly from the nests and runways of their prey, which they have hunted through or taken over. Important prey, e.g. birds (22% of prey items identified) with uninviting nests contributed few fleas (2%); non-prey, e.g. moles (0–4% of items) with highly desirable nests, contributed disproportionately many, including 35 fleas (15%) of two species otherwise monoxenous on moles. A similar pattern is shown by the fleas carried by stoats in New Zealand.     

Phenology and intensity of phyllophage attack on Fagus sylvatica in Wytham Woods, Oxford

ABSTRACT.

• 1 Overwintering beech buds began to swell in early April and bud-break occurred in early May. Most phyllophage attacks were initiated during the 21 day period of leaf growth but additional attacks occurred up to 46 days from bud break.
• 2 An average beech tree, with a top height of 24 m and a d.b.h. of 110 cm, bore 420 345 leaves (17% in the upper canopy, 27% in the upper middle canopy, 27% in the middle canopy, 18% in the lower middle canopy and 11% in the lower canopy).
• 3 Damage by phyllophages was greatest in the lower canopy; some 75—85% of the leaves were attacked in this stratum and damage amounted to 35% of the total phyllopage damage to the trees.
• 4 The major agents of damage in the lower canopy were adult P. argentatus, larval R. fagi and lepidopteran macrophages. Adult R. fagi concentrated their feeding in the upper canopy.
• 5 Allowing for damage expansion due to leaf growth the percentages of the total leaf area actually removed, or seriously damaged, by phyllophages were calculated to be 3.43–4.00% in 1978 and 2.37–2.74% in 1979.

     

Variation of soil and biomass carbon pools in beech forests across a precipitation gradient

Temperate forests have recently been identified as being continuing sinks for carbon even in their mature and senescent stages. However, modeling exercises indicate that a warmer and drier climate as predicted for parts of Central Europe may substantially alter the source/sink function of these economically important ecosystems. In a transect study with 14 mature European beech (Fagus sylvatica L.) forests growing on uniform geological substrate, we analyzed the influence of a large reduction of annual precipitation (970–520 mm yr^?1) on the carbon stocks in fast and slow pools, independent of the well‐known aging effect. We investigated the C storage in the organic L, F, H layers, the mineral soil to 100 cm, and in the biomass (stem, leaves, fine roots), and analyzed the dependence of these pools on precipitation. Soil organic carbon decreased by about 25% from stands with > 900 mm yr^?1 to those with < 600 mm yr^?1; while the carbon storage in beech stems slightly increased. Reduced precipitation affected the biomass C pool in particular in the fine root fraction but much less in the leaf biomass and stem fractions. Fine root turnover increased with a precipitation reduction, even though stand fine root biomass and SOC in the organic L, F, and H layers decreased. According to regression analyses, the C storage in the organic layers was mainly controlled by the size of the fine root C pool suggesting an important role of fine root turnover for the C transfer from tree biomass to the SOC pool. We conclude that the long‐term consequence of a substantial precipitation decrease would be a reduction of the mineral soil and organic layer SOC pools, mainly due to higher decomposition rates. This could turn temperate beech forests into significant carbon sources instead of sinks under global warming.     

Individual biomass factors for beech, oak and pine in Slovakia: a comparative study in young naturally regenerated stands

Biomass conversion and expansion factors (BCEF) which convert tree stem volume to whole tree biomass and biomass allocation patterns in young trees were studied in order to estimate tree and stand biomass in naturally regenerated forests. European beech (Fagus sylvatica L.), Sessile oak (Quercus petraea (Mattuschka) Liebl.) and Scots pine (Pinus sylvestris L.) stands were compared. Seven forest stands of each species were chosen to cover their natural distribution in Slovakia. Species-specific BCEF are presented, generally showing a steep decrease in all species in the smallest trees, with the only exception in the case of branch BCEF in beech which grows with increasing tree size. The values of BCEF for all tree compartments stabilise in all species once trees reach about 60–70-mm diameter at base. As they grow larger, all species increase their allocation to stem and branches, while decreasing the relative growth of roots and foliage. There are, however, clear differences between species and also between broadleaves and conifers in biomass allocation. This research shows that species-specific coefficients must be used if we are to reduce uncertainties in estimates of carbon stock changes by afforestation and reforestation activities.     

Fine root biomass in a beech (Fagus sylvatica L.) stand on Paiko Mountain,NW Greece

Abstract

Fine roots represent a small proportion of total plant biomass however they represent the most dynamic component of the root systems of woody plants. There is limited information on the beech fine root production in Mediterranean ecosystems and especially in Greece. We measured live, dead and total fine root biomass (d<2 mm) (LFRB, DFRB and TFRB, respectively) over a growing season in a beech (Fagus sylvatica L.) stand on Paiko mountain, NW Greece, in order to contribute to the generally scarce knowledge of the fine root biomass of beech stands. It was found that TFRB and LFRB increased from May to July and then decreased. LFRB decreased with soil depth while there was no pattern at the change of DFRB with soil depth.     

Influence of aluminium on biomass, nutrients, soluble carbohydrates and phenols in beech (Fagus sylvatica)

The effects of aluminium on biomass, nutrients and soluble carbohydrates and phenols were studied in beech ( Fagus sylvatica L.) seedlings. After germination, seedlings with cotyledons and the buds of the first leaf-pair developed, were preconditioned for two weeks and then grown for 31 days in nutrient solutions containing 0.1, 0.5, 1.0 or 2.0 m M A1C1₃. Aluminium did not affect the dry weights of roots but at Al concentrations ≥ 1.0 m M the development of the terminal shoot above the first leaf pair, was reduced by 80% or more. The concentrations of most nutrients (P, Ca, Mg, Zn, Cu) in the plant tissues decreased strongly even at the lowest Al levels, but K increased in the shoots. The tissue concentration of N was not affected of Al. but the distribution between the organs was changed to a higher content of N in the roots. At ≥1.0 m M Al the concentrations of starch in both the shoots and the roots were significantly increased, and at ≥ 0.5 m M the roots contained more of total phenols than untreated seedlings. The elevated concentrations and contents of starch and phenols in the seedlings may partly be related to the reduced shoot growth. The observed effects of Al were marked already at Al levels found in soil waters from beech forests in southern Sweden.     

Carabid beetle diversity and mean individual biomass in beech forests of various ages

Carabid beetle diversity and mean individual biomass (MIB) were analysed in three different successional stages of beech tree stands (60, 80 and 150 years old). Carabid beetles were captured using pitfall traps placed at nine sites (three per age class) in the Papuk Mountain of East Croatia during 2008. A cluster analysis identified three groupings that corresponded to the beech age classes. MIB values increased with stand age, ranging from 255 in 60-year-old stand to 537 in the oldest forests. The 80-year-old stand showed the highest species richness and diversity values. With respect to species composition, large species such as Carabus scheidleri and Carabus coriaceus were dominant only in the oldest forests. Furthermore, species that overwinter in the larval stage were more abundant in the oldest forests (45% of the total number of individuals from the 150-year-old stand) than in the younger ones (20% of individuals from 60-year-old, and 22% of individuals from 80-year-old stands). Our results showed that the analyses of species composition and life history traits are valuable for estimating the conservation values of older forests. Although the investigated sites form part of a continuous forested area and are only a couple of kilometres apart, MIB values detect significant differences associated with forest age and can be a useful tool in evaluating the degree to which a forest reflects a natural state.     

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.     

Stand fine root biomass and fine root morphology in old-growth beech forests as a function of precipitation and soil fertility

Only very limited information exists on the plasticity in size and structure of fine root systems, and fine root morphology of mature trees as a function of environmental variation. Six northwest German old-growth beech forests (Fagus sylvatica L.) differing in precipitation (520 – 1030 mm year^–1) and soil acidity/fertility (acidic infertile to basic fertile) were studied by soil coring for stand totals of fine root biomass (0–40 cm plus organic horizons), vertical and horizontal root distribution patterns, the fine root necromass/biomass ratio, and fine root morphology (root specific surface area, root tip frequency, and degree of mycorrhizal infection). Stand total of fine root biomass, and vertical and horizontal fine root distribution patterns were similar in beech stands on acidic infertile and basic fertile soils. In five of six stands, stand fine root biomass ranged between 320 and 470 g m^–2; fine root density showed an exponential decrease with soil depth in all profiles irrespective of soil type. An exceptionally small stand fine root biomass (<150 g m^–2) was found in the driest stand with 520 mm year^–1 of rainfall. In all stands, fine root morphological parameters changed markedly from the topsoil to the lower profile; differences in fine root morphology among the six stands, however, were remarkably small. Two parameters, the necromass/biomass ratio and fine root tip density (tips per soil volume), however, were both much higher in acidic than basic soils. We conclude that variation in soil acidity and fertility only weakly influences fine root system size and morphology of F. sylvatica, but affects root system structure and, probably, fine root mortality. It is hypothesized that high root tip densities in acidic infertile soils compensate for low nutrient supply rates, and large necromasses are a consequence of adverse soil chemical conditions. Data from a literature survey support the view that rainfall is another major environmental factor that influences the stand fine root biomass of F. sylvatica.     

Interactions between fungi,bacteria and beech leaves in a stream microcosm

Summary Immigration and colonization of isolates of naturally occurring stream bacteria and hyphomycetes on beech leaves were studied in a laboratory stream microcosm. Fungal spores were more successful immigrants, especially on new leaves, than bacteria, which were more repelled than attracted by the substrate. Fewer bacteria immigrated to older leaves than to new, and bacteria multiplied faster in water than on leaves. Fungi and bacteria showed synergistic relationships so that each group grew significantly faster in presence of the other group. If one considers, differences in immigration, colonization and synergism patterns, fungal mycelia doubled about 10 times faster than bacterial cells which might explain the dominance of fungi usually found on leaves in early decay. The individual fungal species could be assigned to one of three colonization groups; one of fugitive species, preceding a second group of species that grew from rarity to dominance, and a third group of very slow colonizers. The leachate was fractionated in different molecular size classes by gel chromatography, and the fraction around 2500D in the new leaf leachate was associated with a high concentration of polyphenols. High-pressure liquid chromatography (HPLC) and gas chromatography-mass spectrometry (GC-MS) revealed the presence of 16 phenolic acids in the new leaf leachate at concentrations ranging from < 1 to 640 g l^–1. All fungi except the species with the slowest growth rate grew faster on leachate with the fraction around 2500D removed, and the density of bacteria was significantly reduced when pure stream water was supplemented with compounds from the same fraction.     

Belowground drought response of European beech: fine root biomass and carbon partitioning in 14 mature stands across a precipitation gradient

How tree root systems will respond to increased drought stress, as predicted for parts of Central Europe, is not well understood. According to the optimal partitioning theory, plants should enhance root growth relative to aboveground growth in order to reduce water limitations. We tested this prediction in a transect study with 14 mature forest stands of European beech (Fagus sylvatica L.) by analysing the response of the fine root system to a large decrease in annual precipitation (970–520 mm yr⁻¹). In 3 years with contrasting precipitation regimes, we investigated leaf area and leaf biomass, fine root biomass and necromass (organic layer and mineral soil to 40 cm) and fine root productivity (ingrowth core approach), and analysed the dependence on precipitation, temperature, soil nutrient availability and stand structure. In contrast to the optimal partitioning theory, fine root biomass decreased by about a third from stands with >950 mm yr⁻¹ to those with <550 mm yr⁻¹, while leaf biomass remained constant, resulting in a significant decrease, and not an increase, in the fine root/leaf biomass ratio towards drier sites. Average fine root diameter decreased towards the drier stands, thereby partly compensating for the loss in root biomass and surface area. Both δ¹³C‐signature of fine root mass and the ingrowth core data indicated a higher fine root turnover in the drier stands. Principal components analyses (PCA) and regression analyses revealed a positive influence of precipitation on the profile total of fine root biomass in the 14 stands and a negative one of temperature and plant‐available soil phosphorus. We hypothesize that summer droughts lead to increased fine root mortality, thereby reducing root biomass, but they also stimulate compensatory fine root production in the drier stands. We conclude that the optimal partitioning theory fails to explain the observed decrease in the fine root/leaf biomass ratio, but is supported by the data if carbon allocation to roots is considered, which would account for enhanced root turnover in drier environments.     

Fertilization effects on fineroot biomass, rhizosphere microbes and respiratory fluxes in hardwood forest soils

Fertilizer-induced reductions in CO(2) flux from soil ((F)CO(2)) in forests have previously been attributed to decreased carbon allocation to roots, and decreased decomposition as a result of nitrogen suppression of fungal activity. Here, we present evidence that decreased microbial respiration in the rhizosphere may also contribute to (F)CO(2) reductions in fertilized forest soils. Fertilization reduced (F)CO(2) by 16-19% in 65-yr-old plantations of northern red oak (Quercus rubra) and sugar maple (Acer saccharum), and in a natural 85-yr-old yellow birch (Betula allegheniensis) stand. In oak plots, fertilization had no effects on fine root biomass but reduced mycorrhizal colonization by 18% and microbial respiration by 43%. In maple plots, fertilization reduced root biomass, mycorrhizal colonization and microbial respiration by 22, 16 and 46%, respectively. In birch plots, fertilization reduced microbial respiration by 36%, but had variable effects on root biomass and mycorrhizal colonization. In plots of all three species, fertilization effects on microbial respiration were greater in rhizosphere than in bulk soil, possibly as a result of decreased rhizosphere carbon flux from these species in fertile soils. Because rhizosphere processes may influence nutrient availability and carbon storage in forest ecosystems, future research is needed to better quantify rhizo-microbial contributions to (F)CO(2).     

KlADH3, a gene encoding a mitochondrial alcohol dehydrogenase, affects respiratory metabolism and cytochrome content in Kluyveromyces lactis

A Kluyveromyces lactis strain, harbouring KlADH3 as the unique alcohol dehydrogenase (ADH) gene, was used in a genetic screen on allyl alcohol to isolate mutants deregulated in the expression of this gene. Here we report the characterization of some mutants that lacked or had highly reduced amounts of KlAdh3p activity; in addition, these mutants showed alterations in glucose metabolism, reduced respiration and reduced cytochrome content. Our results confirm that the KlAdh3p activity contributes to the reoxidation of cytosolic NAD(P)H feeding the respiratory chain through KlNdi1p, the mitochondrial internal transdehydrogenase. The low levels of KlAdh3p in two of the mutants were associated with mutations in KlSDH1, one of the genes of complex II, suggesting signalling between the respiratory chain and expression of the KlADH3 gene.     

Modelling temporal decoupling between biomass and numbers during the transient nitrogen-limited growth of a marine phytoflagellate

A mathematical model of single-nutrient-limited algal growthis presented in which carbon fixation and cell division aredifferent functions of assimilated nutrient. The model successfullydescribes key features of the growth of Isochrysis galbana inammonium-limited batch culture under continuous illuminationand in light/dark cycles. The incorporation of a nutrient processingtime allows the simulation of a time lag between net carbonfixation and cell division, and enables the model to describechanges in the mean carbon content and carbon/nitrogen ratioof the cells. The model can be completely parameterized fromstandard batch culture experiments.     

Psychrophilic and psychrotrophic respiratory metabolism in antarctic microplankton

 The activity of the respiratory Electron Transfer System (ETS) was measured in total microplankton (<200-μm size fraction) and nanoplankton (<20-μm size fraction) from the Bransfield Strait, during the ECOANTAR 1993–1994 cruise of the Spanish B.I.O. Hespérides in January 1994. Activity variation in response to temperature was measured at three stations belonging to three different water masses that showed in situ temperatures ranging from −0.57 to 1.30^°C. Subsamples from each station were assayed for ETS activity at 11 temperatures in the −3 to 20^°C range. The results showed a bimodal activity-temperature variation in plankton from the lower in situ temperatures, with a peak in activity at 0^°C, and a minimum at 3^°C, with subsequent continuous increase up to absolute maxima at 15^°C. The water mass with higher than 0^°C temperature did not show the 0^°C activity peak. The results suggest the existence, in water masses with in situ temperature near or below 0^°C, of psychrophilic microbial populations with a narrow temperature range of respiratory enzyme activity, coexisting with more numerous and widespread psychrotrophs, or cold-tolerant populations, whose ETSs showed a continuous increase in activity in the −3 to 15^°C temperature range. Arrhenius activation energies (E_a) of total microplankton ranged from 3 to 17 kcal mole^-1, and the Q₁₀ from 1.2 to 3.5. These facts point to the existence of differentiated biochemical adaptations and acclimations to low temperature in polar plankton, an issue that has been much discussed in the recent past. Received: 20 July 1995/Accepted: 28 October 1995