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.     

The effect of plant species,weather variables and chemical composition of plant material on decomposition in a tropical grassland

Summary The decomposition of litter and roots ofChenopodium album, Desmostachya bipinnata and mixed grass samples for a period of 402 days and ofDichanthium annulatum andSesbania bispinosa for a period of 278 days was studied in a tropical grassland. Litter bags positioned at midcanopy height, soil surface and at five cm depth below the soil surface and root bags placed at 5, 15, 25 and 35 cm depths belowground were used. For the total study period, the cumulative weight loss in litter bags was: Chenopodium=76–100%; Desmostachya=33–98%; Dichanthium=26–96%; mixed grass=43–99% and Sesbania=25–99%. The weight loss in root bags was: Chenopodium=93–100%; Desmostachya=47–56%; Dichanthium=71–87%; mixed grass=61–82%; Sesbania=87–100%. The nature of plant species affected decomposition rates. The position of litter/root bags also affected the decomposition rates. The mean relative decomposition rates of litter as well as of root material were found to be highest in rainy season and lowest in winter months. Rainfall, particularly the frequency of rainfall, was an important factor affecting decomposition rates. The litter species characterized by highest concentration of nitrogen, ash, acid detergent cell wall component and lowest concentration of carbon, cellulose and lignin, decomposed rapidly. In the case of roots, the material having high nitrogen, carbon, cellulose and ash content and low C/N ratio and lignin content decomposed rapidly.     

Intraspecific variability in leaf traits strongly affects alder leaf decomposition in a stream

This study assessed the intraspecific variability of senescent leaves of alder (Alnus glutinosa Gaertn.) and the effects of this variability on leaf decomposition in streams. Leaves were collected at five geographically distant locations in Europe. We analyzed 10 batches of leaf samples for seven quantitative leaf traits as well as leaf decomposition rate in coarse and fine mesh bags exposed in a single stream. The geographic origin of leaf samples largely explained the observed variation in litter quality and decomposition rate. Phosphorus (0.034–0.187%) and lignin (3.9–18.7%) concentrations in leaves varied widely. Together, these two traits accurately predicted leaf decomposition rate (r²=84.1%). Intraspecific variation in leaf decomposition rate was within a range similar to that reported for interspecific variation among co-occurring riparian plant species in Europe. Our study demonstrates extensive intraspecific variability in leaf traits on a continental scale, which can have enormous effects on major ecosystem processes such as leaf decomposition.     

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.     

Rapid nutrient cycling in leaf litter from invasive plants in Hawai’i

Physiological traits that contribute to the establishment and spread of invasive plant species could also have impacts on ecosystem processes. The traits prevalent in many invasive plants, such as high specific leaf areas, rapid growth rates, and elevated leaf nutrient concentrations, improve litter quality and should increase rates of decomposition and nutrient cycling. To test for these ecosystem impacts, we measured initial leaf litter properties, decomposition rates, and nutrient dynamics in 11 understory plants from the Hawaiian islands in control and nitrogen + phosphorus fertilized plots. These included five common native species, four of which were ferns, and six aggressive invasive species, including five angiosperms and one fern. We found a 50-fold variation in leaf litter decay rates, with natives decaying at rates of 0.2–2.3 year^–1 and invaders at 1.4–9.3 year^–1. This difference was driven by very low decomposition rates in native fern litter. Fertilization significantly increased the decay rates of leaf litter from two native and two invasive species. Most invasive litter types lost nitrogen and phosphorus more rapidly and in larger quantities than comparable native litter types. All litter types except three native ferns lost nitrogen after 100 days of decomposition, and all litter types except the most recalcitrant native ferns lost >50% of initial phosphorus by the end of the experiment (204–735 days). If invasive understory plants displace native species, nutrient cycling rates could increase dramatically due to rapid decomposition and nutrient release from invasive litter. Such changes are likely to cause a positive feedback to invasion in Hawaii because many invasive plants thrive on nutrient-rich soils.     

Forest floor biomass,litter fall and nutrient return in Central Himalayan oak forests

The seasonal dynamics of forest floor biomass, pattern of litter fall and nutrient return in Central Himalayan oak forests are described. Fresh and partially decomposed litter layers occur throughout the whole year in addition to herbaceous vegetation. The highest leaf litter value is found in April and May and the minimum in September. Partially and largely decomposed litter tended to increase from January to May with a slight decline in June. The wood litter peaked in March and April. The relative contribution of partially decomposed litter to the forest floor remains greatest the year round. The maximum herbaceous vegetation development was found in September with a total annual net production of 104.3 g m^-2yr^-1. The total calculated input of litter was 480.8 g m^-2yr^-1. About 68% of the forest floor was replaced each year with a subsequent turnover time of 1.47 yr. The total annual input of litter ranged from 664 (Quercus floribunda site) –952 g m^-2 (Q. lanuginosa site), of which tree, shrub and herbaceous litter accounted for respectively 72.0–86.3%, 6.4 – 19.4% and 5.2 – 8.6%. The annual nutrient return through litter fall amounted to (kg ha^-1) 178.0 – 291.0 N, 10.0 – 26.9 P, 176.8 – 301.6 Ca, 43.9 – 64.1 K and 3.98 – 6.45 Na. The tree litter showed an annual replacement of 66.0 – 70.0%, for different nutrients the range was 64 and 84%.     

凋落物多样性及组成对凋落物分解及土壤微生物群落的影响——二氧化碳倍增条件下

受全球变化的影响生物多样性的丧失日益严重,及时了解凋落物物种多样性及其组成如何直接或者通过调节微生物群落来间接影响凋落物分解已经成为生态学领域的热点问题之一。在呼伦贝尔典型草原区,羊草（Leymus chinensis）为原生群落建群种,茵陈蒿（Artemisia capillaris）、麻花头（Serratula centauroides）、二裂委陵菜（Potentilla bifurca）在退化及恢复群落中的多度均较大,本研究以此4种植物的凋落物为研究对象,在两倍于当前大气CO₂浓度（800 μmol/mol）条件下,通过嵌套实验设计来研究凋落物多样性（凋落物组成）对凋落物质量、C、N残余率和残余C/N的影响,并深入分析凋落物初始性质以及土壤革兰氏阳性菌（G⁺）、革兰氏阴性菌（G^-）、细菌（B）、真菌（F）及土壤总微生物磷脂脂肪酸（Phospholipid Fatty Acid,PLFA）含量和F/B对凋落物分解的影响。结果表明：（1）凋落物多样性及组成对凋落物质量、C、N残余率以及残余C/N均具有显著影响;凋落物组成对G⁺ PLFAs含量具有显著影响;（2）冗余分析（Redundancy Analysis,RDA）结果表明凋落物组成对凋落物分解和土壤微生物群落相关指标的影响高于凋落物多样性;（3）结构方程模型（Structural Equation Model,SEM）结果表明凋落物初始木质素含量和初始C/N均对凋落物分解具有显著影响,其中凋落物初始木质素含量起主导作用,其还可通过对土壤真菌PLFAs含量的影响来间接影响凋落物N残余率和残余C/N。所得结果可为大气CO₂浓度升高条件下退化草原生态系统的物质循环特征的预测乃至草原生态系统功能的合理评估提供数据支持。     

Litter quality in a north European transect versus carbon storage potential

Newly shed foliar plant litter often has a decomposition rate of ca 0.1–0.2% day^–1, which decreases greatly with time and may reach 0.0001 to 0.00001% day^–1 or lower in litter material in the last stages of decay. The decrease in decomposability (substrate quality) varies among species and is complex, involving both direct chemical changes in the substrate itself and the succession in microorganisms able to compete for substrate with a given chemical composition. In late stages, the decomposition appears very little affected by climate, suggesting that climate change will have little effect on late-stages decomposition rates. Here, we apply a model for the late stages of litter decomposition to address the question of climate-change effects on soil-C storage. Decomposition of litter turning into soil organic matter (SOM) is determined by the degradation rate of lignin. In the last phases of decay, raised N concentrations have a rate-retarding effect on lignin degradation and thus on the decomposition of far-decomposed litter and litter in near-humus stages. The retardation of the decomposition rate in late stages may be so strong that decomposition reaches a limit value at which total mass losses virtually stop. At such a stage the remaining litter would be close to that of stabilized SOM. The estimated limit values for different species range from about 45 to 100% decomposition indicating that between 0 and 55% should either be stabilized or decompose extremely slowly. For no less than 106 long-term studies on litter decomposition, encompassing 21 litter types, limit values were significantly and negatively related to N concentration, meaning that the higher the N concentration in the newly shed litter (the lower the C/N ratio) the more litter was left when it reached its limit value. Trees growing under warmer and wetter climates (higher actual evapotranspiration, AET) tend to shed foliar litter more rich in N than those growing under colder and drier climates. A change in climate resulting in higher AET would thus mean that within species, e.g., Scots pine, a higher N level in the foliar litter may result. Further, within the boreal system deciduous species appear to have foliar litter richer in N than have conifers and within the conifers group, Norway spruce has needle litter more rich in N than, e.g., Scots pine. Thus, a change of species (e.g., by planting) from pine to spruce or from spruce to a deciduous species such as birch may result in a higher N level in the litter fall at a given site. In both cases the result would be a lower limit value for decomposition. The paper presents an hypothesis, largely based on available data that a change in climate of 4° higher annual average temperature and 40% higher precipitation in the Baltic basin would result in higher N levels in litter, lower decomposition and thus a considerable increase in humus accumulation.     

Mineral cycling in a tropical palm forest

Nutrient cycling and biomass characteristics of a tropical palm forest dominated byOrbignya cohune were found to be different from thsoe of hardwood dominated forests. The cohune palm forest had a high proportion of biomass in leaves (5%), a reduced sapling layer, a large amount of standing forest litter and an exceptionally low decomposition rate factor (0.1 year^–1). Mineral concentrations in palm leaves were generally lower than in hardwood species with the exception of Na, which was exceptionally high inOrbignya cohune. Biomass was estimated at 226 tons ha^–1 containing 1173 kg ha^–1 N; 126 kg ha^–1 P; 437 kg ha^–1 K; 1869 kg ha^–1 Mg; 125 kg ha^–1 Ca, and 2177 kg ha^–1 Na. Soils of cohune association did not differ significantly from those of neighbouring hardwood dominated associations with the exception of Na which occurred in higher concentration because of bioaccumulation in the dominant. The results suggest that the growth habits and physiology of a dominant can strongly influence some of the ecological parameters used to describe aforest association.     

Leaf litter decomposition of Piper aduncum,Gliricidia sepium and Imperata cylindrica in the humid lowlands of Papua New Guinea

No information is available on the decomposition and nutrient release pattern of Piper aduncum and Imperata cylindrica despite their importance in shifting cultivation systems of Papua New Guinea and other tropical regions. We conducted a litter bag study (24 weeks) on a Typic Eutropepts in the humid lowlands to assess the rate of decomposition of Piper aduncum, Imperata cylindrica and Gliricidia sepium leaves under sweet potato (Ipomoea batatas). Decomposition rates of piper leaf litter were fastest followed closely by gliricidia, and both lost 50% of the leaf biomass within 10 weeks. Imperata leaf litter decomposed much slower and half-life values exceeded the period of observation. The decomposition patterns were best explained by the lignin plus polyphenol over N ratio which was lowest for piper (4.3) and highest for imperata (24.7). Gliricidia leaf litter released 79 kg N ha^–1, whereas 18 kg N ha^–1 was immobilised in the imperata litter. The mineralization of P was similar for the three species, but piper litter released large amounts of K. The decomposition and nutrient release patterns had significant effects on the soil. The soil contained significantly more water in the previous imperata plots at 13 weeks due to the relative slow decomposition of the leaves. Soil N levels were significantly reduced in the previous imperata plots due to immobilisation of N. Levels of exchangeable K were significantly increased in the previous piper plots due to the large addition of K. It can be concluded that piper leaf litter is a significant and easily decomposable source of K which is an important nutrient for sweet potato. Gliricidia leaf litter contained much N, whereas imperata leaf litter releases relatively little nutrients and keeps the soil more moist. Gliricidia fallow is more attractive than an imperata fallow for it improves the soil fertility and produces fuelwood as additional saleable products.     

Factors affecting nitrogen dynamics in a semiarid woodland (Dry Chaco,Argentina)

In an effort to elucidate the factors affecting soil N dynamics in the Dry Chaco ecosystem, soil respiration and microbial biomass N were measured for one year underneath 5 vegetation types: a leguminous tree (Prosopis flexuosa DC), a non-leguminous tree (Aspidosperma quebracho-blanco Schlecht.), a non leguminous shrub (Larrea spp.), the open interspaces, and a pure grassland. Ammonifier and nitrifier densities and N content in litter were also measured in some cases. Results were compared with previously reported N mineralization rates and soil fertility.During the dry season microbial biomass N and net N mineralization were low, while accretion of easily mineralizable C occurred (estimated through soil respiration rates in lab under controlled temperature and moisture). With the onset of rain, microbial biomass N and N mineralization increased markedly, resulting in a decrease in easily mineralizable C. Throughout the wet season N mineralization varied with soil moisture while microbial biomass N remained consistently high. Mean values of immobilized N in this ecosystem were high (20–140 mg kg^–1), of about the same order of magnitude as accumulated net N mineralization (50–150 mg kg^–1 yr^–1). Microbial decay in the dry season, considered as a source of easily mineralizable N, accounted for only 40% of gross N mineralization increase at the beginning of the wet season. Ammonifier densities correlated significantly with soil moisture and N mineralization, but nitrifiers did not.The highest values of total N, N mineralization, inorganic N, microbial biomass N, nitrifier densities, N content in litter, total organic C and easily mineralizable C were found under Prosopis and the lowest values under shrubs and the interspaces. The main differences between tree species were in N mineralization at the beginning of the wet season, in total and inorganic N pools, and in nitrifier densities; all of which were significantly lower under Aspidosperma than under Prosopis.N mineralization in the pure grassland was very low despite high values of total N and C sources. Although N immobilized in microbial biomass was similarly high under Aspidosperma, Prosopis and the pure grassland, net N mineralization rates were quite different.     

Litter decomposition contrasts in second- and old-growth Douglas-fir forests of the Pacific Northwest, USA

Litterfall and its subsequent decomposition are important feedback mechanisms in the intrasystem cycling of nutrients in forest ecosystems. The amount of litterfall and the rate of decomposition are expected to vary with stand age and climate. Over a 2-year period, decomposition of five litter types were measured in two second-growth forest stands and one old-growth stand in the Cascade Mountains of southern Washington state, USA. Both second-growth stands were dominated by Douglas-fir [Pseudotsuga menziesii (Mirb.,) Franco] but one had a significant proportion of red alder (Alnus rubra Bong.), a nitrogen (N) fixer. The old-growth stand was dominated by Douglas-fir and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. All stands had a relatively shallow layer of forest floor mass. The five litter types were placed in each stand to evaluate decomposition patterns. Despite significant differences in stand age, microclimate and mean residence times for carbon (C) and N, the rates of litter mass loss varied only slightly between sites. The relative order of species litter mass loss was: vine maple ≫ salal = western hemlock > Douglas-fir (from the youngest stand) > Douglas-fir (from the N rich stand with red alder). The initial litter lignin concentration, not lignin:N, was the primary determinant of decomposition rates, although the initial N concentration was the predictor for mass loss after 2 years in the N rich Douglas-fir-alder stand. All litter types showed immobilization of N for nearly 2 years. Data for Douglas-fir litter suggest that higher levels of N may retard decomposition of tissues with greater amounts of lignified material. The retention of N by the litter appeared influenced by the nutrient capital of the stands as well as the forest floor C:N ratio. Decomposition was minimal during the cold winter months, but displayed a definitive peak period during early Fall with wet weather, warm soils, and fungal activity. Thus, long-term climatic change effects on forest floor C storage may depend more on changes in seasonality of precipitation changes than just temperature changes.     

Nutrient limitation to the decomposition of water hyacinth ( Eichhornia crassipes)

The responses of decomposition to N and P supply were investigated in three leaf types of water hyacinth (Eichhornia crassipes (Mart.) Solms): dead green leaves collected from Donghu Lake; green, and brown leaves collected from outdoor tanks. The ratios of C:N, C:P, lignin:N and lignin:P were lowest in the green leaves collected from Donghu Lake, and highest in the brown leaves collected from outdoor tanks. Decomposition constant (k) of water hyacinth varied greatly, ranged from 0.006 to 0.099 d^–1. Leaf litters decayed most quickly within the initial two weeks during the experimental period, but decomposition rate decreased significantly in the following days. Decomposition and nutrient (N and P) release were fastest in the green leaves collected from Donghu Lake, intermediate in the green leaves collected from outdoor tanks, slowest in the brown leaves collected from outdoor tanks. Statistical analyses revealed that the effects of P-availability on decomposition rate and N, P release rate of the three litter types were significant, whereas the impacts of N-availability was insignificant (p > 0.05) except for the brown leaves collected from outdoor tanks. These results suggest that decomposition rate and nutrient content dynamics of water hyacinth differ with their growth habitats, and could partly be regulated by nutrient availability, especially by P-availability, in the environments.     

Decomposition of macrophyte litter in a subtropical constructed wetland in south Florida (USA)

The South Florida Water Management District has constructed large treatment wetlands (stormwater treatment areas (STAs)) to reduce total phosphorus concentrations in agricultural runoff before this water enters the Everglades. An important component of nutrient removal and storage in these systems is incorporation of nutrients into aquatic macrophytes and burial of this biomass in the sediments. However, decomposition of plant biomass before burial returns nutrients to the water column and may reduce STA treatment efficiency. As part of research on biogeochemical control of STA performance, we conducted a summer (July–September) and a long-term (12-month) experiment (February–February) that measured decomposition rates and release of chemical constituents from dominant aquatic macrophytes in a constructed wetland located in south Florida. The rank order of mean decomposition rates was Najas/Ceratophyllum (0.0568 d⁻¹) > Pistia (0.0508 d⁻¹) > Eichhornia (0.0191 d⁻¹) > submerged Typha (0.0059 d⁻¹) > aerial Typha (0.0008 d⁻¹). Summer decomposition rates were generally higher than rates from the long-term experiment, which suggested a temperature effect. Decomposition rates were negatively correlated with litter C:N and C:P molar ratios and cellulose and lignin content and positively correlated with N and P content. There was no significant difference in decomposition rates among sampling stations despite the fact that there was a decreasing gradient in water column inorganic phosphorus and nitrogen concentrations at these sites. Relatively little of the initial P mass remained in the litter of all species, except Typha, by the end of both experiments. First-order decomposition models derived using nonlinear regression generally had explanatory power, i.e. accounted for variance, comparable to more complex decreasing-coefficient models. Decomposition rates for the species examined in this study were within the range of published values when comparisons were made either by species or by plant group.     

漓江流域岩溶区檵木群落不同恢复阶段凋落物分解初期动态

为探究漓江流域岩溶区檵木群落不同恢复阶段凋落物的分解情况,运用凋落物袋法研究其凋落物分解初期动态。结果表明:经过1a的分解,檵木群落凋落物失重大小顺序为:灌木阶段乔灌阶段乔林阶段。檵木群落灌木阶段、乔灌阶段和乔林阶段凋落物分解50%所需的时间分别为1.28a、1.38a、1.41a,分解95%所需的时间分别为5.54a、5.97a和6.09a。经过1a的分解,凋落物养分动态变化为:灌木阶段和乔灌阶段C含量总体上升,乔林阶段C含量总体下降;3个恢复阶段N和纤维素含量总体上升;灌木阶段和乔林阶段P含量总体下降,乔灌阶段P含量总体上升;灌木阶段木质素含量总体上升,而乔灌阶段和乔林阶段木质素含量总体下降。相关分析表明,灌木阶段凋落物分解速率分别与C,N,P,C/P,N/P之间呈显著或极显著相关性,乔灌阶段凋落物分解速率与N和木质素/N之间呈显著或极显著相关性,乔林阶段凋落物分解速率分别与N,纤维素、C/N和木质素/N之间呈显著或极显著相关性。     

Litter quality,decomposition rates and saprotrophic mycoflora in Fallopia japonica (Houtt.) Ronse Decraene and in adjacent native grassland vegetation

Fallopia japonica succeeds in invading different ecosystems likely because of its huge biomass production. This biomass is characterized by low nutritional quality and low decomposition rates but knowledge on whether these features are correlated to microbial decomposers is still lacking. The aims of this work were: i) to determine litter decomposition rates of native grassland vegetation and F. japonica under different conditions in a year-round experiment; ii) to evaluate litter quality and/or site effect on the decomposition of the invader and native vegetation and iii) to characterize mycoflora isolated from F. japonica and native vegetation litter. The results showed that F. japonica litter decomposes 3–4 times slower than that of native grassland, mainly due to its low N content and consequently high C/N ratio both in leaves and stems. As decomposition proceeds C/N in F. japonica litter decreases to values approaching those of the grassland litter. Site had no effect on the decomposition rates of F. japonica and grassland litter. Total fungal load and composition differed between F. japonica and native litter, and also varied across sites. These results indicate that the successful invasive plant F. japonica affects the structure and functions of the invaded ecosystem through a huge production of low quality, slow-decomposing litter that selects saprotrophic fungi.     

Decomposition patterns of leaf litter of seven common canopy species in a subtropical forest: N and P dynamics

Litter decomposition, governing nutrient and C cycling, is strongly influenced by the chemical litter quality. In order to determine the interspecific variation in leaf decomposition rates and to understand the chemical basis for such variation, decomposition dynamics of seven common canopy species was investigated over 2year using the litterbag technique in a subtropical evergreen broad-leaved forest on Okinawa Island, Japan. The species studied are representatives of the vegetation in the study area and differed significantly in their chemical litter quality. Dry mass loss at the end of study varied in the order: Distylium racemosum< Quercus miyagii< Rapanea neriifolia< Symplocos confusa< Castanopsis sieboldii< Schima wallichii< Daphniphyllum glaucescens. All species showed a pattern characterized by a rapid initial decomposition followed by lower rates except for D. glaucescenswhich decomposition rate appeared to be rather constant. In the late phase, decomposition rates were correlated positively to initial N and ash contents and negatively to lignin content, lignin:N, C:N, and C:P ratios. The effects of N and lignin content or lignin:N ratio were stronger than other quality parameters. There was a wide range in patterns of N and P concentrations, from a net accumulation to a rapid loss in decomposition. The correlation between N and P release suggests that N and P dynamics may have influenced each other during litter decomposition. Analysis of initial quality for species showed that the C:P ratios were extremely high (range 1639–3811) but the N:P ratios were from 28 to 56, indicating a likely P-limitation for this forest. Our results suggest that P is an important control of litter decomposition and N and P dynamics.     

Factors affecting formation of cluster roots in Myrica gale seedlings in water culture

The effect of nutrient deficiency, aeration, phosphorus supply, and nitrogen source on the formation of cluster (proteoid) roots was examined in Myrica gale seedlings growing in water culture. Only the omission of phosphorus resulted in the formation of significant numbers to cluster roots when plants were grown in a number of 1/4 strength Hoagland's solutions, each lacking one mineral nutrient. Aeration shortened the time required for cluster root formation and increased the percentage of plants forming cluster roots. The proportion of the root system comprised of cluster roots decreased as the phosphorus concentration in the solution increased and no cluster roots formed in solutions containing 8 mg P/L. Phosphorus supply also affected total plant biomass, proportion of biomass comprising nitrogen-fixing nodules, shoot:root ratio, phosphorus concentration in the leaves and phosphorus content of the plants. The plants showed luxury consumption of phosphorus and were able to produce large amounts of biomass utilizing only stored phosphorus.Nitrogen source also affected cluster root formation. Urea-fed plants produced cluster roots more quickly and devoted a substantially larger proportion of root growth to cluster roots than did nitrate-fed plants. The longest cluster root axes were produced in nitrate-fed plants supplied with no phosphorus and the shortest were in urea-fed plants at 4 mg P L^–1.Four methods for expressing the extent of cluster root formation were examined and it was concluded that cluster roots as a proportion of total fine root dry weight is preferable in many cases. Formation of cluster roots in response to phosphorus deficiency coupled with previously demonstrated traits allows Myrica gale to adapt to a wide range of soil conditions.     

Decay rate and substrate quality of fine roots and foliage of two tropical tree species in the Luquillo Experimental Forest,Puerto Rico

Decomposition rates, initial chemical composition, and the relationship between initial chemistry and mass loss of fine roots and foliage were determined for two woody tropical species, Prestoea montana and Dacryodes excelsa, over a gradient of sites in two watersheds in the Luquillo Experimental Forest, Puerto Rico. At all locations, fine roots decayed significantly more slowly than foliage during the initial 6 months.Substrate quality of the initial tissue showed marked differences between roots and foliage when using cell wall chemistry, secondary chemistry and total elemental analysis as indices. Quantity of acid detergent fiber (ADF) (non-digestible cell wall fiber) and lignin content were higher for roots than leaves: D. excelsa roots had 55.3% ADF and 28.7% lignin while leaves had 36.2% ADF and 11.8% lignin; P. montana roots had 68.0% ADF and 26.8% lignin while leaves had 48.5% ADF and 16.1% lignin. Aluminum concentrations were higher in fine roots (843 mg kg^–1 in D. excelsa, 1500 mg kg^–1 in P. montana) than leaves (244 mg kg^–1 in D. excelsa, 422 mg kg^–1 in P. montana), while calcium concentrations were higher in foliage (5.5 mg g^–1 in D. excelsa, 7.8 mg g^–1 in P. montana) than roots (3.4 mg g^–1 in D. excelsa, 3.1 mg g^–1 in P. montana). Nitrogen did not show any trend with tissue or species type. A linear model between mass remaining after 6 months and initial tissue chemistry could be developed only for calcium (r²=0.64).     

Non‐symbiotic nitrogen fixation during leaf litter decomposition in an old‐growth temperate rain forest of Chiloé Island,southern Chile: Effects of single versus mixed species litter

Heterotrophic nitrogen fixation is a key ecosystem process in unpolluted, temperate old‐growth forests of southern South America as a source of new nitrogen to ecosystems. Decomposing leaf litter is an energy‐rich substrate that favours the occurrence of this energy demanding process. Following the niche ‘complementarity hypothesis’, we expected that decomposing leaf litter of a single tree species would support lower rates of non‐symbiotic N fixation than mixed species litter taken from the forest floor. To test this hypothesis we measured acetylene reduction activity in the decomposing monospecific litter of three evergreen tree species (litter C/N ratios, 50–79) in an old‐growth rain forest of Chiloé Island, southern Chile. Results showed a significant effect of species and month (anova , Tukey's test, P < 0.05) on decomposition and acetylene reduction rates (ARR), and a species effect on C/N ratios and initial % N of decomposing leaf litter. The lowest litter quality was that of Nothofagus nitida (C/N ratio = 78.7, lignin % = 59.27 ± 4.09), which resulted in higher rates of acetylene reduction activity (mean = 34.09 ± SE = 10.34 nmol h^?1 g^?1) and a higher decomposition rate (k = 0.47) than Podocarpus nubigena (C/N = 54.4, lignin % = 40.31 ± 6.86, Mean ARR = 4.11 ± 0.71 nmol h^?1 g^?1, k = 0.29), and Drimys winteri (C/N = 50.6, lignin % = 45.49 ± 6.28, ARR = 10.2 ± 4.01 nmol h^?1 g^?1, k = 0.29), and mixed species litter (C/N = 60.7, ARR = 8.89 ± 2.13 nmol h^?1g^?1). We interpret these results as follows: in N‐poor litter and high lignin content of leaves (e.g. N. nitida) free‐living N fixers would be at competitive advantage over non‐fixers, thereby becoming more active. Lower ARR in mixed litter can be a consequence of a lower litter C/N ratio compared with single species litter. We also found a strong coupling between in situ acetylene reduction and net N mineralization in surface soils, suggesting that as soon N is fixed by diazotroph bacteria it may be immediately incorporated into mineral soil by N mineralizers, thus reducing N immobilization.