Foliage litter turnover and earthworm populations in three beech forests of contrasting soil and vegetation types

Summary Leaf litter decomposition, levels of accumulated litter as well as the abundance and biomass of earthworms were measured in three mature beech forests in southern Sweden: one mor site, one poor mull site, and one rich mull site. The disappearance rate of beech litter, measured with litter bags, increased with increasing soil fertility. On the rich mull site, the disappearance rate was much higher than in the two other forests, due to the combined effects of higher earthworm activity, more favouable soil moisture conditions, and higher litter quality. Incubating the litter in finely meshed bags (1-mm mesh) to exclude macrofauna had a great effect on litter mass loss in the rich mull site, but it had only a minor effect in the other sites. Simultaneous incubations of local and transplanted leaf litter on the three study sites showed that the substrate quality of the litter increased in the order: mor site — poor mull site — rich mull site. Lignin, N, and P concentrations of the leaf litter failed to explain the observed differences in decomposition rates, and acid/base properties are suggested to be more important. Earthworm numbers per m² were 2.5 (1 species) in the mor, 40 (6 species) in the poor mull and 220 (9 species) in the rich mull forest. Soil chemical conditions, notably pH, were suggested as the main factors determining the inter-site differences in abundance and species composition of earthworms. The role of litter decomposition and earthworm activity in the accumulation of organic matter in the forest floor in different types of beech woodlands are discussed.     

The spatial pattern and canopy-understory association of trees in a cool temperate,mixed forest in western Japan

The spatial distribution pattern of trees and the association between canopy and understory individuals were examined with reference to the distribution of tree crowns in a cool temperate, mixed forest in Ohdaigahara, western Japan. Line transect and contact sampling methods were used to examine the pattern over various spatial scales. These methods are useful to detect patterns over a large study area. The dominance ofChamaecyparis obtusa on steep slopes forming large patches suggested that the distribution of this species is a consequence of landslides. UnderstoryFagus crenata showed a clumped distribution, and the relative coverage of this species was larger in canopy gaps than under a closed canopy. Understory individuals ofAbies homolepis showed a positive association with canopy trees ofF. crenata but a negative association with conspecific canopy individuals. These patterns suggested thatF. crenata regenerates in canopy gaps and is replaced byA. homolepis. The dynamics of these two species are consistent with the process of gap dynamics. The effects of both small- and large-scale disturbance must be evaluated to understand the mechanisms of patch formation and the coexistence of forest tree species.     

A 5 year study of litter decomposition processes in aChamaecyparis obtusa Endl. forest

Decomposition of needle litter in aChamaecyparis obtusa forest was studied over a 5 year period using a litter bag method. Organic matter, nitrogen and carbon mass and faunal abundance were monitored. The pattern of weight loss was represented by three phases: the initial leaching of carbon and nitrogen (0–3 months), nitrogen immobilization (3–15 months), and nitrogen mobilization (15–60 months). The decomposition rate of needle litter was expressed by Olson's decomposition constant (k) which was−0.113 over a 5 year period. The decomposition rate decreased with the advance of decomposition processes. The role of soil fauna in the decomposition process was assessed by comparing decomposition rates between the control and defaunated plots. In the leaching phase, soil animals had no significant role in the decomposition processes. During the immobilization phase, soil animals contributed to the immobilization processes through their grazing activities, and there were significant differences in weight loss between the control and defaunated plots. In the mobilization phase, saprovorous soil animals such as Collembola and Acari contributed to the mobilization processes by feeding on decomposing litter. Decomposition processes observed in this study were compared with other similar studies.     

Effect of legume understorey on decomposition and nutrient content of eucalypt forest litter

Summary In Jarrah (Eucalyptus marginata Donn ex Sm.) forest of south-western Australia dense germination and regeneration of the native legumeAcacia Pulchella R. Br. can occur following moderate to high intensity fire. The effect of this legume understorey on rate of decomposition and change in nutrient content ofE. marginata litter was investigated using the mesh bag techniques and by examining four components of forest floor litter representing increasing stages of decomposition. E. marginata leaf litter confined in mesh bags lost 37% of its initial dry weight in the first 8 months on the forest floor and 44% of its initial dry weight after 20 months. During this period weight loss was similar for leaf litter located in forest without legume understorey and for leaf litter placed under dense stands ofA. pulchella. MixingA. pulchella litter withE. marginata litter had no significant effect on rate ofE. marginata litter breakdown. The presence of understorey vegetation had a marked effect on chemical composition of decomposingE. marginata leaves. After 8 and 20 months exposure on the forest floor, leaf litter in mesh bags placed underA. pulchella understorey had significantly (P<0.001) higher concentration and contained significantly (P<0.001) greater amounts of N, P, K, S, Ca and Mg than leaf litter placed in areas without legume understorey. This effect was particularly marked for N and P. In forest without legume understorey the amounts of these two nutrients inE. marginata leaf litter changed little during the first 20 months of decomposition, but forE. marginata leaf litter in mesh bags underA. pulchella there were absolute gains of up to 68% in the amount of N and 109% in the amount of P during this period. This represents accumulation of N and P from sources outside the litter bags. The concentration of N, P, S, Ca and Mg were higher at each of the four stages of decomposition in eucalypt leaf litter collected from the forest floor beneathA. pulchella compared to eucalypt leaf litter collected in forest without understorey. Concentrations of N, P and S increased with stage of decomposition. Levels of these three nutrients in eucalypt litter from under the legume were 1.5 to 2.9 fold higher than in the same component of litter from forest without understorey. The effect of legume understorey on nutrient concentrations in the forest floor and on Cielement ratios in decomposing litter is discussed in relation to long term rates of litter breakdown and net mineralisation of litter nutrients.     

Leaf litter quality and decomposition rates of yellow birch and sugar maple seedlings grown in mono-culture and mixed-culture pots at three soil fertility levels

Seedlings of yellow birch (Betula alleghaniensis Britton) and sugar maple (Acer saccharum Marsh.) were grown for 2 years in mono-culture and mixed-culture and at three fertility levels. Following the second growing season, senescent leaves were analysed for N concentration, acid hydrolysable substances (AHS), and nonhydrolysable remains (NHR). A litter sub-sample was then inoculated with indigenous soil microflora, incubated 14 weeks, and mass loss was measured. Litter-N was significantly higher at medium than at poor fertility, as well as in yellow birch than in sugar maple litter. The species effect on litter-N increased with increasing fertility. At medium fertility, litter-N of sugar maple litter was lower in mixed-culture than in mono-culture. AHS, NHR as well the NHR/N ratio were significantly higher in yellow birch than in sugar maple litter. At medium fertility, the NHR/N ratio of sugar maple litter was significantly lower in mono-culture than in mixed-culture. Mass loss was significantly greater at medium and rich fertility than at poor fertility, and in yellow birch than in sugar maple litter. At poor fertility, mixed-litter decomposed at a rate comparable to yellow birch, whereas at medium and rich fertility, mixed-litter decomposed at a rate comparable to sugar maple. There was a significant positive relationship between litter-N and mass loss. A similar positive relationship between NHR and mass loss was presumed to be a species effect on decomposition. Results support the hypothesis that species × fertility and species × mixture interactions can be important determinants of litter quality and, by implication, of site nutrient cycling.     

Seedling recruitment ofChamaecyparis obtusa andSciadopitys verticillata in different microenvironments in an old-growthSciadopitys verticillata forest

Seedling recruitment ofChamecyparis obtusa andSciadopitys verticillata in different microenvironments in an old-growthS. verticillata forest was studied for five years. Exposed mineral soil beneath a gap, which was newly created by a typhoon, supported the best seedling emergence forC. obtusa. S. verticillata seemed to require no specific microenvironment for its seedling emergence. Mortality of seedlings was higher for both species beneath a closed canopy, where all new seedlings ofC. obtusa died within a year after their emergence. Beneath the gap, mortality of seedlings was generally lower for both species, regardless of the presence of litter on the ground. The occurrence of saplings was more frequent forC. obtusa beneath the gaps and forS. verticillata beneath a closed canopy. Difference of seedling recruitment behaviour could maintain the coexistence ofC. obtusa andS. verticillata in the canopy layer of this forest in a non-equilibrium state caused by occasional disturbance.     

Somatic embryogenesis and plant regeneration from immature zygotic embryos of Hinoki cypress (Chamaecyparis obtusa Sieb. et Zucc.)

We established a plant regeneration system for Hinoki cypress (Chamaecyparis obtusa) via somatic embryogenesis. Embryogenic tissues were successfully induced on three kinds of Smith media from megagametophyte explants containing pre-cotyledonary embryos of C. obtusa plus-trees. Factors affecting somatic embryo maturation were examined. The concentration of polyethylene glycol 4000 in the medium was a critical factor for embryo maturation and its effective concentration was 150 g/l. The addition of 30 g/l maltose to the medium had a positive effect on embryo maturation, but sucrose was ineffective. The mature somatic embryos germinated at a germination frequency of approximately 60%, and the presence of activated charcoal was effective in stimulating plantlet growth. The plantlets acclimatized successfully in a greenhouse. To our knowledge, this is first report describing details of a plant regeneration method for C. obtusa via somatic embryogenesis.Abbreviations ABA Abscisic acid - PEG Polyethylene glycol 4000 - SM1 Smith Standard Embryonic Tissue Capture Medium - SM2 Smith Standard Embryogenesis Medium - SM3 Smith Embryo Develop Medium     

Leaf litter decomposition and nutrient dynamics in a subtropical forest after typhoon disturbance

Decomposition of typhoon-generated and normal leaf litter and their release patterns for eight nutrient elements were investigated over 3 yr using the litterbag technique in a subtropical evergreen broad-leaved forest on Okinawa Island, Japan. Two common tree species, Castanopsis sieboldii and Schima wallichii, representative of the vegetation and differing in their foliar traits, were selected. The elements analyzed were N, P, K, Ca, Mg, Na, Al, Fe and Mn. Dry mass loss at the end of study varied in the order: typhoon green leaves > typhoon yellow leaves > normal leaves falling for both species. For the same litter type, Schima decomposed faster than Castanopsis. Dry mass remaining after 2 yr of decomposition was positively correlated with initial C:N and C:P ratios. There was a wide range in patterns of nutrient concentration, from a net accumulation to a rapid loss in decomposition. Leaf litter generated by typhoons decomposed more rapidly than did the normal litter, with rapid losses for N and P. Analysis of initial quality for the different litter types showed that the C:P ratios were extremely high (range 896 – 2467) but the P:N ratios were < 0.05 (range 0.02 – 0.04), indicating a likely P-limitation for this forest. On average 32% less N and 60% less P was retranslocated from the typhoon-generated green leaves than from the normal litter for the two species, Castanopsis and Schima. An estimated 2.13 g m^–2 yr^–1 more N and 0.07 g m^–2 yr^–1 more P was transferred to the soil as result of typhoon disturbances, which were as high as 52% of N and 74% of P inputted from leaf litter annually in a normal year. Typhoon-driven maintenance of rapid P cycling appears to be an important mechanism by which growth of this Okinawan subtropical forest is maintained.     

Immunolocalization of glucomannans in the cell wall of differentiating tracheids inChamaecyparis obtusa

Summary A polyclonal antibody against glucomannans (GMs) was raised in a mouse. A dot-blot immunoassay and competitive-inhibition tests indicated that the antibody was specific for GMs. The antibody enables visualization of the localization of GMs in differentiating tracheids ofChamaecyparis obtusa. Labeling of GMs was restricted to the secondary walls of the tracheids. The labeling density temporarily increased and then decreased in the outer and middle layers of the secondary wall during cell wall formation. This is probably due to the accumulation of lignin. In comparison with previous studies of glucuronoxylans, there must be a clear difference between the deposition of GMs and that of glucuronoxylans.     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Tree leaf litter composition and nonnative earthworms influence plant invasion in experimental forest floor mesocosms

Dominant tree species influence community and ecosystem components through the quantity and quality of their litter. Effects of litter may be modified by activity of ecosystem engineers such as earthworms. We examined the interacting effects of forest litter type and earthworm presence on invasibility of plants into forest floor environments using a greenhouse mesocosm experiment. We crossed five litter treatments mimicking historic and predicted changes in dominant tree composition with a treatment of either the absence or presence of nonnative earthworms. We measured mass loss of each litter type and growth of a model nonnative plant species (Festuca arundinacea, fescue) sown into each mesocosm. Mass loss was greater for litter of tree species characterized by lower C:N ratios. Earthworms enhanced litter mass loss, but only for species with lower C:N, leading to a significant litter × earthworm interaction. Fescue biomass was significantly greater in treatments with litter of low C:N and greater mass loss, suggesting that rapid decomposition of forest litter may be more favorable to understory plant invasions. Earthworms were expected to enhance invasion by increasing mass loss and removing the physical barrier of litter. However, earthworms typically reduced invasion success but not under invasive tree litter where the presence of earthworms facilitated invasion success compared to other litter treatments where earthworms were present. We conclude that past and predicted future shifts in dominant tree species may influence forest understory invasibility. The presence of nonnative earthworms may either suppress of facilitate invasibility depending on the species of dominant overstory tree species and the litter layers they produce.     

Identification of calmodulin in the green alga Mougeotia and its possible function in chloroplast reorientational movement

A soluble protein was isolated from Mougeotia by chloropromazine-sepharose 4 B affinity chromatography. The protein matches the properties of calmodulin in terms of heat stability, Ca²⁺-dependent electrophoretic mobility in sodium-dodecyl-sulfate polyacrylamide gels, and its ability to activate cyclic nucleotide phosphodiesterase in a Ca²⁺-dependent manner. Phytochrome-mediated chloroplast reorientational movement in Mougeotia was inhibited by the calmodulin antagonist trifluoperazine, a hydrophobic compound, or N-(6-aminohexyl)-5-chloro-1-naphthalenesulfonamide (W-7), a hydrophilic compound; 50% inhibition (IC₅₀) of chloroplast movement is caused by 20–50 mol l^-1 trifluoperazine or 100 mol l^-1 W-7. The Ca²⁺-calmodulin may act as an intermediate in the chloroplast reorientational response in Mougeotia governed by phytochrome.Abbreviations EGTA ethylene glycol-bis(-aminoethyl ether)-N,N,N,N-tetraacetic acid - SDS sodium dodecyl sulfate - W-7 N-(6-aminohexyl)-5-chloro-1-naphthalene sulfonamide     

Fine root growth and nutrient release in decomposing leaf litter in three contrasting vegetation types in central Amazonia

We tested the hypothesis that the growth of fine roots upward into the leaf litter, forming a ‘surface root mat’, found widely in Amazonian forests, is of adaptive value for plants of poor soils because it makes possible more rapid uptake of limiting nutrients. We assessed the effect of invasion by fine roots on the rates of loss of dry mass and nutrient content of leaf litter over 1 year in three plots in each of three contrasting forest types in central Amazonia: the stunted facies of heath forest known as campina (SHF), the taller facies of heath forest known as campinarana (THF), both on spodosols, and the surrounding lowland evergreen rain forest (LERF) on ultisol. Pairs of bags filled with freshly fallen leaves from the trees of Clitoria fairchildiana (Fabaceae) were placed on the litter layer; in order to prevent roots entering the control bag in each pair that bag was lifted from the forest floor and turned over each week, while the treatment bag was left undisturbed. From each plot, four pairs of litter bags were retrieved after 30, 60, 120, 180, 270 and 360 d, and all roots that had grown into the litterbags were carefully removed. The leaves and roots from each bag were oven-dried for nutrient analysis. In no forest type was there a significant difference in the rate of loss of dry matter from the litter between the bags with and without roots. The amounts of roots invading the litter bags increased significantly in the sequence SHF < THF < LERF. In no forest did the presence of roots have a significant effect on the rate of disappearance of N or P from the leaf material, or on the rate of accumulation of Fe and Al. In the SHF there was no significant effect of invasion by roots on the rates of disappearance of Ca, Mg, Mn or Zn, but in the THF, the rates of disappearance of these four elements between 270 and 360 d were significantly greater in the presence of roots. In the LERF the results were similar, but the effects of roots started earlier—the rates of disappearance of Ca and Mg were significantly enhanced between 120 and 360 d. The results from the SHF may be interpreted to suggest that the growth of fine roots (and their fungal associates) upward into leaf litter is unlikely to be explained wholly by their role in the efficient recovery of mineral nutrients.     

Concentrations of lignin and wall-bound ferulic acid after wounding in the phloem of Chamaecyparis obtusa

The changes of lignin and wall-bound ferulic acid induced by wounding were quantitatively and histochemically investigated in the phloem of Chamaecyparis obtusa. Histochemical staining of lignin was first observed in the necrotic region of the phloem 7 days after wounding and developed in 14 days. Increases of the wall-bound ferulic acid and lignin concentrations were detected in the necrotic tissue at 7 and 14 days, respectively. The concentrations continued to increase until 28 days. The lignin concentration of the callus tissue was observed to be lower than that of the healthy tissue at 14 days, and reached a similar level after 28 days. No quantitative changes of lignin and wall-bound ferulic acid were observed in other tissues. The results indicated that lignin synthesis could be maintained after the phloem cells were discolored and seemed to be necrotic. Distribution and timing of the wall-bound ferulic acid and lignin accumulation suggested that the increased wall-bound ferulic acid was involved in the lignin synthesis after wounding.     

Agrobacterium tumefaciens-mediated transformation of embryogenic tissue and transgenic plant regeneration in Chamaecyparis obtusa Sieb. et Zucc.

A genetic transformation procedure for Chamaecyparis obtusa was developed after co-cultivation of embryogenic tissues with disarmed Agrobacterium tumefaciens strain C58/pMP90, which harbours the sgfp (synthetic green fluorescent protein) visual reporter and nptII (neomycin phoshotransferase II) selectable marker genes. The highest transformation frequency was 22.5 independent transformed lines per dish (250 mg embryogenic tissue) following selection on kanamycin medium. Transgenic plantlets were regenerated through the maturation and germination of somatic embryos. The intensity of GFP fluorescence, observed under a fluorescence microscope, varied from very faint to relatively strong, depending on the transgenic line or part of the transgenic plant. The integration of the genes into the genome of regenerated plantlets was confirmed by Southern blot analysis.     

Analytical studies on migrating movement of the pseudo-plasmodium ofDictyostelium discoideum

Summary Migrating movement of a pseudoplasmodium (slug) of the cellular slime mouldDictyostelium discoideum was analyzed using a time-lapse video tape recorder. Since slugs usually migrated with repeated interruptions of advance, migrating velocities were measured only within a period of forward movement. On the basis of some known facts and assumptions, a dynamical model for slug movement was formulated, which consists of motive force generated by slug cells against their intrinsic resistance and resistance of slime sheath at the tip. The migrating velocity of a slug depended neither on its width nor its volume, but solely on its length. Under any experimental conditions tested, a linear relationship always held between reciprocals of the two variables. The results were in good agreement with predictions of the model. Quantitative analyses of experimental results by the use of the model lead to the conclusions that a decrease in velocity at a low temperature is due to an increase in resistance of slime sheath at the tip, but that a decrease in velocity during prolonged migration is due to a decrease in motive force of constituent cells. An anterior isolate dissected from a slug migrated at a velocity greater than that of an intact slug of the same length. This was interpreted by the model to be due to the fact that the anterior cells have greater motive forces and intrinsic resistances than the posterior cells. The heterogeneous distributions of the two variables in the cell mass is discussed in reference to the mechanism of sorting out of cells.     

Chemical composition of litter affects the growth and enzyme production by the saprotrophic basidiomycete Hypholoma fasciculare

Chemical composition of litter has previously been reported to affect in situ decomposition. To identify its effects on a single species level, the saprotrophic basidiomycete Hypholoma fasciculare was grown on 11 types of litter with variable chemical composition (N content of 3.4–28.9 mg g⁻¹), and the mass loss of litter and lignin, production of extracellular enzymes and fungal biomass were followed. After 12 weeks, mass loss ranged from 16 % to 34 %. During early decomposition stages, litter mass loss, fungal biomass production (estimated by ergosterol content) as well as fungal substrate use efficiency all increased with increasing initial N content of the litter. The initial litter decomposition rate was significantly positively correlated with the activities of arylsulfatase, cellobiohydrolase, endoxylanase and phosphatase. Contrary to expectations, the lignin content did not affect litter mass loss, when covariation with N content was accounted for. The ratio of lignin loss to total mass loss depended on the litter type and did not reflect the activities of ligninolytic enzymes.     

Stand development and regeneration during a 33-year period in a seral Picea glehnii forest, northern Japan

Stand development and regeneration were studied during a 33-year period (1965-1998) in a 1-ha plot in a seral Picea glehnii forest in northern Japan. P. glehnii was mono-dominant in the upper canopy layer, but its understory trees were rarely found in 1965. Other species were scarcely observed in 1965. Many recruited saplings of Abies sachalinensis which had grown to > 5 cm diameter at breast height (DBH) by 1998 had become dominant in the understory layer. Mortality of P. glehnii canopy trees was low. Therefore, the stand basal area increased during the census period due to the growth of surviving canopy trees. Stand development brought about intense competition among trees by increasing local crowding for each tree, and promoted dominance of larger trees and suppression of smaller trees. Although growth rates of understory trees of the two conifers decreased with the increase in local crowding, the growth rate of A. sachalinensis was consistently higher than that of P. glehnii at all extents of local crowding. The recruitment rate (growing to 5 cm DBH) of the two conifers was less affected by local crowding. However, the number of recruits of P. glehnii was only about a quarter of that of A. sachalinensis during the census period because the regeneration of P. glehnii was largely restricted to fallen logs and within 1 m of the base of any live tree > 20 cm DBH. Therefore, our long-term study suggests that A. sachalinensis will dominate over P. glehnii in the seral forest because of higher recruitment and growth rates of the former than the latter in the understory.An erratum to this article can be found at     

The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance

Summary We studied the effect of removing and adding plant litter in different seasons on biomass, density, and species richness in a Solidago dominated old-field community in New Jersey, USA. We removed all the naturally accumulated plant litter in November (658 g/m²) and in May (856 g/m²) and doubled the amount of litter in November and May in replicated plots (1 m²). An equal number of plots were left as controls. Litter removal and addition had little impact on total plant biomass or individual species biomass in the growing season following the manipulations. Litter removal, however, significantly increased plant densities but this varied depending upon the season of litter removal, species, and life history type. Specifically, the fall litter removal had a much greater impact than the spring litter removal suggesting that litter has its greatest impact after plant senescence in the fall and prior to major periods of early plant growth in spring. Annual species showed the greatest response, especially early in the growing season. Both spring and fall litter removal significantly increased species richness throughout the study. Litter additions in both spring and fall reduced both plant densities and species richness in June, but these differences disappeared near the end of the growing season in September. We concluded than in productive communities where litter accumulation may be substantial, litter may promote low species richness and plant density. This explanation does not invoke resource competition for the decline in species richness. Finally, we hypothesize that there may be broad thresholds of litter accumulation in different community types that may act to either increase or decrease plant yield and diversity.