Convergence and divergence of nutrient stoichiometry during forest litter decomposition

Background and aims

General theory of forest floor dynamics indicates convergence of properties during detrital decomposition. This study examined the hypothesis that nutrient stoichiometry, i.e. the relative amounts of nutrients, converges during litter decomposition.

Methods

Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and red alder (Alnus rubra Bong.) foliar litters that decomposed in their respective forests were analyzed for N, P, Ca, Mg, K, S, Mn, Fe, Zn, and Cu. A novel approach quantified the stoichiometric difference as the angle between nutrient vectors. The approach was also used to synthesize data from 11 previously published studies representing a broad array of litter types and locations.

Results

The stoichiometries of the Douglas-fir and red alder litters converged during the first 2 years of decomposition, but diverged in the subsequent 4 years. This temporal trajectory was explained by two competing sets of processes: stoichiometric convergence occurs when different litters decompose in the same environment, and divergence occurs when the same litter decomposes in different environments. Manganese, Fe, and Ca were important contributors to stoichiometric differences.

Conclusions

Stoichiometric convergence processes often dominate over divergence processes. Consideration of multi-nutrient stoichiometries may enhance the understanding of the functions of litter, including rates of decomposition and relative rates at which nutrients are released.     

Fate and effects of phosphorus additions in soils under N2-fixing red alder

Soil phosphorus (P) dynamics are controlled by the interaction of geochemical, biochemical and biological processes. Changes in species composition or management could alter the relative importance of these processes. We examined soil P dynamics in two plantations of N₂-fixing red alder (Alnus rubra) by determining the fate and effects of added fertilizer P. History of the plantations varied such that sites were previously occupied by 60-yr-old stands of alder or non-fixing Douglas-fir (Pseudotsuga menziesii). Without fertilization, the soil with a longer period of alder influence had more organic P (P_o) and less sorbed inorganic P (Hydroxide- and Bicarb-extractable P_i). Fertilization increased soil total P, and 88% of the fertilizer was accounted for in the surface mineral soil (0–15 cm). Sorbed P_i was the major sink for fertilizer P (55–60%), independent of site history. Although P_o was 35–70% of soil P in unfertilized plots, added P did not accumulate as P_o. Neither site history nor P addition influenced phosphatase activity. Fertilization increased decomposition during incubation of the organic horizon, suggesting that late-stage decomposition is P-limited in these N-rich soils. On the time-scale of a few years, geochemical sorption and desorption of inorganic P were the most important processes controlling the distribution of added P. Organic P accumulation is expected to occur over a longer time frame, linked to the production and turnover of organic matter.     

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.     

Influence of stand age on nutrient and energy release through decomposition in alder-cardamom agroforestry systems of the eastern Himalayas

The influence of stand age (5, 10, 15, 20, 30 and 40 years) on the decomposition of litter fractions, nutrient and energy release of mixtures of N₂-fixing alder (Alnus nepalensis) and non-N₂-fixing large cardamom (Amomum subulatum) systems was compared. Seasonal decomposition rates were distinct with the highest rate in the first 6 months followed by subsequent seasons. The decomposition rate was substantially high in younger stands (10- to 15-years) and declined in the older stands. Heat sink from the stand floor litter increased from 171 × 10⁶ kJ year⁻¹ in 5 years to 299 × 10⁶ kJ year⁻¹ at 15 years and then considerably decreased with advancing age. However, energy and nutrient releases were slow at a high initial lignin-to-initial N ratio and C-to-N ratio, and there was an inverse relationship between the k-value of ash-free-mass and N expressed as a function of the C-to-N ratio. Quantities of nutrient release and energy loss per unit area in 24 months of decomposition were highest in 15 years and subsequently they lowered with advancing age. Nutrient loss indicated approximately uniform absolute and relative rates. Absolute energy consistently decreased by 81–88% in 24 months. Ash-free mass of decomposing litter remaining at different retrieval dates was associated with a narrowing of the C-to-N ratio. The relative loss rate of ash-free mass, nutrients and energy content was strongly related to the C-to-N ratio, litter temperature and litter moisture. The influence of Alnus in the younger stands on nutrient and energy releases were rapid, indicating accelerated nutrient cycling and energy dynamics. The intensity of the processes was highly phenomenal and considerably high in younger stands up to 20 years. Thus, an appropriate management cycle of the Alnus-cardamom system for sustainability is 15–20 years.     

Growth and N2 fixation in an Alnus hirsuta (Turcz.) var. sibirica stand in Japan

To estimate the N₂ fixation ability of the alder (Alnus hirsuta (Turcz.) var. sibirica), we examined the seasonal variation in nitrogenase activity of nodules using the acetylene reduction method in an 18-year-old stand naturally regenerated after disturbance by road construction in Japan. To evaluate the contribution of N₂ fixation to the nitrogen (N) economy in this alder stand, we also measured the phenology of the alder, the litterfall, the decomposition rate of the leaf litter, and N accumulation in the soil. The acetylene reduction activity per unit nodule mass (ARA) under field conditions appeared after bud break, peaked the maximum in midsummer after full expansion of the leaves, and disappeared after all leaves had fallen. There was no consistent correlation between ARA and tree size (dbh). The amount of N₂ fixed in this alder stand was estimated at 56.4 kg ha^?1 year^?1 when a theoretical molar ratio of 3 was used to convert the amount of reduced acetylene to the amount of fixed N₂. This amount of N₂ fixation corresponded to the 66.4% of N in the leaf litter produced in a year. These results suggested that N₂ fixation still contributed to the large portion of N economy in this alder stand.     

Impacts of Falcataria moluccana Invasion on Decomposition in Hawaiian Lowland Wet Forests: The Importance of Stand-level Controls

Invasive species have the capacity to substantially alter soil processes, including rates of litter decomposition. Currently, the few remaining native-dominated lowland wet forests in Hawai’i are being invaded by Falcataria moluccana, a large, fast-growing, N₂-fixing tree. In this study, we sought to determine the extent to which Falcataria invasion alters decomposition in these lowland wet forests, and whether changes resulted from differences in litter substrate type, lava flow age and type, forest stand type and associated soil biota, or some combination of these factors. We measured decomposition rates and nitrogen (N) and phosphorus (P) dynamics of Metrosideros polymorpha and Falcataria leaf litter in native-dominated and Falcataria-invaded stands on 48- and 300-year-old a’a lava flows and a 213-year-old pāhoehoe flow in the Puna district of eastern Hawai’i. Despite significant differences in the initial quality of Metrosideros and Falcataria litter, in nearly all cases mass remaining of the two litter types did not differ within a given forest stand, whether native-dominated or invaded. Instead, stand type accounted for large differences in the decomposition of both litter types, and litter decomposed two to 10 times faster in Falcataria-invaded stands than it did in their native-dominated counterparts on each lava flow. Dynamics of N (that is, immobilization or release) during decomposition were affected by stand, litter, and lava flow type; P dynamics were affected by stand and flow type, but not litter type. Although not definitive proof of causality, the decay rates of both species were positively correlated to previously measured inputs of N mass and P mass via litterfall as well as availability of soil N and P, characteristics that all increased substantially with Falcataria invasion. Given the degree of change to a host of ecosystem processes, including decomposition, after invasion by Falcataria, these transformed forest ecosystems may best be viewed as fundamentally new and different, in both structure and function, from the native ecosystems they have replaced.     

Elevated temperature may intensify the positive effects of nutrients on microbial decomposition in streams

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A study of nodulation and nitrogen fixation of alder on the purplish soils in China

Summary The alder has a perennial nodule cluster. The nodule amount on the roots increases with tree age. The N₂-fixing activity of nodules decreases with nodule age. Purple coloured soils with various soil pHs and CaCO₃ contents are, in the main, the ones which influence nodulation and N₂-fixing. Higher N₂-fixing capacity existed in the neutral and low calcium soils. High calcium soils and acid soils can restrain nodulation and the N₂-fixing rate significantly. On the slope, where calcarous light loams are found, the annual nitrogen fixation capacity of alder and cypress mixed plantations, less than 10 years old, is 16 or 17 kg/ha yr, but in the valley, a pure alder plantation can reach 40 kg/ha yr.     

Benefit to N<Subscript>2</Subscript>-fixing alder of extending growth period at the cost of leaf nitrogen loss without resorption

This study examines the adaptive role of not resorbing N in N₂-fixing deciduous trees in terms of their energy balance. The autumnal growth of N₂-fixing Alnus firma Sieb. et Zucc. (alder) was compared with that of the non-N₂-fixing Morus bombycis Koizumi (mulberry), which resorbs leaf N. The freezing resistance of leaves of both species was –2°C. Mulberry seedlings lost their photosynthetic ability in mid-October, although the minimum temperature was still above 0°C. Thereafter, their leaves turned yellow and were gradually shed. In contrast, seedlings of the alder maintained their photosynthetic ability until mid-November, when the minimum temperature fell to the freezing resistance limit. Thereafter, their leaves were shed quickly without an autumn tint. The mulberry resorbed 48.9% of leaf N, whereas the alder resorbed hardly any. These results show that, compared with the mulberry tree, the alder extended its growth period for 1 month in return for losing leaf N without resorption. The amount of energy assimilated by the alder in the extended growth period was about six times that required for compensating for the nitrogen loss, if the compensation is dependent only on the tree's own nitrogen fixation. This surplus energy balance has probably allowed N₂-fixing deciduous trees to evolve their non-N-resorbing trait.     

Does nitrogen availability control rates of litter decomposition in forests?

The effects of increased exogenous N availability on rates of litter decomposition were assessed in several field fertilization trials. In a jack pine (Pinus banksiana Lamb.) forest, needle litter decomposed at the same rate in control plots and in plots fertilized with urea and ammonium nitrate (1350 kg N ha^-1) with or without P and K. Mixed needle litter of western hemlock (Tsuga heterophylla (Raf.) Sarg.), western red cedar (Thuja plicata Donn) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) incubated in plots recently amended with sewage sludge (500 kg N ha^-1) lost less weight during 3 years than did litter in control plots. Forest floor material also decomposed more slowly in plots amended with sewage sludge. Paper birch (Betula papyrifera Marsh.) leaf litter placed on sewage sludge (1000 kg N ha^-1), pulp sludge, or sewage-pulp sludge mixtures decomposed at the same rate as leaf litter in control plots. These experiments demonstrate little effect of exogenous N availability on rates of litter decomposition.The influence of endogenous N availability on rates of litter decomposition was examined in a microcosm experiment. Lodgepole pine (Pinus contorta var. latifolia Engelm.) needle litter collected from N-fertilized trees (525 kg N ha^-1 in ammonium nitrate) were 5 times richer in N than needles from control trees (1.56% N versus 0.33% N in control trees), but decomposed at the same rate. Green needles from fertilized trees contained twice as much N as needles from control trees (1.91% N versus 0.88% N), but decomposed at the same rate. These experiments suggest that N availability alone, either exogenous or endogenous, does not control rates of litter decomposition. Increased N availability, through fertilization or deposition, in the absence of changes in vegetation composition, will not alter rates of litter decomposition in forests.     

Alder and lupine enhance nitrogen cycling in a degraded forest soil in Northern Sweden

Positive effects of legumes and actinorhizal plants on N-poor soils have been observed in many studies but few have been done at high latitudes, which was the location of our study. We measured N₂ fixation and several indices of soil N at a site near the Arctic Circle in northern Sweden. More than 20 years ago lupine (Lupinus nootkatensis Donn) and gray alder (Alnus incana L. Moench) were planted on this degraded forest site. We measured total soil N, net N mineralization and nitrification with a buried bag technique, and fluxes of NH⁺ ₄ and NO^– ₃ as collected on ion exchange membranes. We also estimated N₂ fixation activity of the N₂-fixing plants by the natural abundance of ¹⁵N of leaves with Betula pendula Roth. as reference species. Foliar nitrogen in the N₂-fixing plants was almost totally derived from N₂ fixation. Plots containing N₂-fixing species generally had significantly higher soil N and N availability than a control plot without N₂-fixing plants. Taken together, all measurements indicated that N₂-fixing plants can be used to effectively improve soil fertility at high latitudes in northern Sweden.     

Nitrogen availability affects saprotrophic basidiomycetes decomposing pine needles in a long term laboratory study

Fungi, especially basidiomycetous litter decomposers, are pivotal to the turnover of soil organic matter in forest soils. Many litter decomposing fungi have a well-developed capacity to translocate resources in their mycelia, a feature that may significantly affect carbon (C) and nitrogen (N) dynamics in decomposing litter. In an eight-month long laboratory study we investigated how the external availability of N affected the decomposition of Scots pine needles, fungal biomass production, N retention and N-mineralization by two litter decomposing fungi – Marasmius androsaceus and Mycena epipterygia. Glycine additions had a general, positive effect on fungal biomass production and increased accumulated needle mass loss after 8 months, suggesting that low N availability may limit fungal growth and activity in decomposing pine litter. Changes in the needle N pool reflected the dynamics of the fungal mycelium. During late decomposition stages, redistribution of mycelium and N out from the decomposed needles was observed for M. epipterygia, suggesting autophagous self degradation.     

Elevated CO2 increases the long-term decomposition rate of Quercus myrtifolia leaf litter

Decomposition of Quercus myrtifolia leaf litter in a Florida scrub oak community was followed for 3 years in two separate experiments. In the first experiment, we examined the effects CO₂ and herbivore damage on litter quality and subsequent decomposition. Undamaged, chewed and mined litter generated under ambient and elevated (ambient+350 ppm V) CO₂ was allowed to decompose under ambient conditions for 3 years. Initial litter chemistry indicated that CO₂ levels had minor effects on litter quality. Litter damaged by leaf miners had higher initial concentrations of condensed tannins and nitrogen (N) and lower concentrations of hemicellulose and C : N ratios compared with undamaged and chewed litter. Despite variation in litter quality associated with CO₂, herbivory, and their interaction, there was no subsequent effect on rates of decomposition under ambient atmospheric conditions. In the second experiment, we examined the effects of source (ambient and elevated) of litter and decomposition site (ambient and elevated) on litter decomposition and N dynamics. Litter was not separated by damage type. The litter from both elevated and ambient CO₂ was then decomposed in both elevated and ambient CO₂ chambers. Initial litter chemistry indicated that concentrations of carbon (C), hemicellulose, and lignin were higher in litter from elevated than ambient CO₂ chambers. Despite differences in C and fiber concentrations, litter from ambient and elevated CO₂ decomposed at comparable rates. However, the atmosphere in which the decomposition took place resulted in significant differences in rates of decomposition. Litter decomposing under elevated CO₂ decomposed more rapidly than litter under ambient CO₂, and exhibited higher rates of mineral N accumulation. The results suggest that the atmospheric conditions during the decomposition process have a greater impact on rates of decomposition and N cycling than do the atmospheric conditions under which the foliage was produced.     

Cascading effects of induced terrestrial plant defences on aquatic and terrestrial ecosystem function

Herbivores induce plants to undergo diverse processes that minimize costs to the plant, such as producing defences to deter herbivory or reallocating limited resources to inaccessible portions of the plant. Yet most plant tissue is consumed by decomposers, not herbivores, and these defensive processes aimed to deter herbivores may alter plant tissue even after detachment from the plant. All consumers value nutrients, but plants also require these nutrients for primary functions and defensive processes. We experimentally simulated herbivory with and without nutrient additions on red alder (Alnus rubra), which supplies the majority of leaf litter for many rivers in western North America. Simulated herbivory induced a defence response with cascading effects: terrestrial herbivores and aquatic decomposers fed less on leaves from stressed trees. This effect was context dependent: leaves from fertilized-only trees decomposed most rapidly while leaves from fertilized trees receiving the herbivory treatment decomposed least, suggesting plants funnelled a nutritionally valuable resource into enhanced defence. One component of the defence response was a decrease in leaf nitrogen leading to elevated carbon : nitrogen. Aquatic decomposers prefer leaves naturally low in C : N and this altered nutrient profile largely explains the lower rate of aquatic decomposition. Furthermore, terrestrial soil decomposers were unaffected by either treatment but did show a preference for local and nitrogen-rich leaves. Our study illustrates the ecological implications of terrestrial herbivory and these findings demonstrate that the effects of selection caused by terrestrial herbivory in one ecosystem can indirectly shape the structure of other ecosystems through ecological fluxes across boundaries.     

Decomposition nitrogen is better retained than simulated deposition from mineral amendments in a temperate forest

Nitrogen (N) deposition (N_DEP) drives forest carbon (C) sequestration but the size of this effect is still uncertain. In the field, an estimate of these effects can be obtained by applying mineral N fertilizers over the soil or forest canopy. A ¹⁵N label in the fertilizer can be then used to trace the movement of the added N into ecosystem pools and deduce a C effect. However, N recycling via litter decomposition provides most of the nutrition for trees, even under heavy N_DEP inputs. If this recycled litter nitrogen is retained in ecosystem pools differently to added mineral N, then estimates of the effects of N_DEP on the relative change in C (?C/?N) based on short‐term isotope‐labelled mineral fertilizer additions should be questioned. We used ¹⁵N labelled litter to track decomposed N in the soil system (litter, soils, microbes, and roots) over 18 months in a Sitka spruce plantation and directly compared the fate of this ¹⁵N to an equivalent amount in simulated N_DEP treatments. By the end of the experiment, three times as much ¹⁵N was retained in the O and A soil layers when N was derived from litter decomposition than from mineral N additions (60% and 20%, respectively), primarily because of increased recovery in the O layer. Roots expressed slightly more ¹⁵N tracer from litter decomposition than from simulated mineral N_DEP (7.5% and 4.5%) and compared to soil recovery, expressed proportionally more ¹⁵N in the A layer than the O layer, potentially indicating uptake of organic N from decomposition. These results suggest effects of N_DEP on forest ?C/?N may not be apparent from mineral ¹⁵N tracer experiments alone. Given the importance of N recycling, an important but underestimated effect of N_DEP is its influence on the rate of N release from litter.     

Nitrogen Export from Forested Watersheds in the Oregon Coast Range: The Role of N<Subscript>2</Subscript>-fixing Red Alder

Variations in plant community composition across the landscape can influence nutrient retention and loss at the watershed scale. A striking example of plant species importance is the influence of N₂-fixing red alder (Alnus rubra) on nutrient cycling in the forests of the Pacific Northwest. To understand the influence of red alder on watershed nutrient export, we studied the chemistry of 26 small watershed streams within the Salmon River basin of the Oregon Coast Range. Nitrate and dissolved organic nitrogen (DON) concentrations were positively related to broadleaf cover (dominated by red alder: 94% of basal area), particularly when near-coastal sites were excluded (r ² = 0.65 and 0.68 for nitrate-N and DON, respectively). Nitrate and DON concentrations were more strongly related to broadleaf cover within entire watersheds than broadleaf cover within the riparian area alone, which indicates that leaching from upland alder stands plays an important role in watershed nitrogen (N) export. Nitrate dominated over DON in hydrologic export (92% of total dissolved N), and nitrate and DON concentrations were strongly correlated. Annual N export was highly variable among watersheds (2.4–30.8 kg N ha^–1 y^–1), described by a multiple linear regression combining broadleaf and mixed broadleaf–conifer cover (r² = 0.74). Base cation concentrations were positively related to nitrate concentrations, which suggests that nitrate leaching increases cation losses. Our findings provide evidence for strong control of ecosystem function by a single plant species, where leaching from N saturated red alder stands is a major control on N export from these coastal watersheds.     

Litterfall and litter accumulation in red alder stands in western Oregon

Summary Litter production in red alder communities 2 to 33 years old was higher than that reported for any plant community of the temperate regions. Litter production is also atypically high in communities of other nitrogen fixers, which suggests that nitrogen fixation may be an important contribution to the high litter production. Litter accumulates rapidly on the ground during the first 5 years after red alder establishment, but equilibrium is reached at the age of 6 years and maintained for several decades. During this time, the decomposition of litter equals the annual litterfall. In 50 years, cumulative litterfall reaches over 300 metric tons per ha, most of which is decomposed and incorporated into mineral soil. Favorable physical, chemical, and nutritional conditions of soil are created by alder for development of the climax vegetation. Paper No.733.     

Insect biological control accelerates leaf litter decomposition and alters short‐term nutrient dynamics in a Tamarix‐invaded riparian ecosystem

Insect herbivory can strongly influence ecosystem nutrient dynamics, yet the indirect effects of herbivore‐altered litter quality on subsequent decomposition remain poorly understood. The northern tamarisk beetle Diorhabda carinulata was released across several western states as a biological control agent to reduce the extent of the invasive tree Tamarix spp. in highly‐valued riparian ecosystems; however, very little is currently known about the effects of this biocontrol effort on ecosystem nutrient cycling. In this study, we examined alterations to nutrient dynamics resulting from beetle herbivory in a Tamarix‐invaded riparian ecosystem in the Great Basin Desert in northern Nevada, USA, by measuring changes in litter quality and decomposition, as well as changes in litter quantity. Generally, herbivory resulted in improved leaf litter chemical quality, including significantly increased nitrogen (N) and phosphorus (P) concentrations and decreased carbon (C) to nitrogen (C:N), C:P, N:P, and lignin:N ratios. Beetle‐affected litter decomposed 23% faster than control litter, and released 16% more N and 60% more P during six months of decomposition, as compared to control litter. Both litter types showed a net release of N and P during decomposition. In addition, herbivory resulted in significant increases in annual rates of total aboveground litter and leaf litter production of 82% and 71%, respectively, under the Tamarix canopy. Our finding that increased rates of N and P release linked with an increased rate of mass loss during decomposition resulting from herbivore‐induced increases in litter quality provides new support to the nutrient acceleration hypothesis. Moreover, results of this study demonstrate that the introduction of the northern tamarisk beetle as biological control to a Tamarix‐invaded riparian ecosystem has lead to short‐term stimulation of nutrient cycling. Alterations to nutrient dynamics could have implications for future plant community composition, and thus the potential for restoration of Tamarix‐invaded ecosystems.     

Effects of elevated CO<Subscript>2</Subscript> and nitrogen on wood structure related to water transport in seedlings of two deciduous broad-leaved tree species

Wood structure might be altered through the physiological responses to atmospheric carbon dioxide concentration ([CO₂]) and nitrogen (N) deposition. We investigated growth, water relations and wood structure of 1-year-old seedlings of two deciduous broad-leaved tree species, Quercus mongolica (oak, a ring-porous species) and Alnus hirsuta (alder, a diffuse-porous species and N₂–fixer), grown under a factorial combination of two levels of [CO₂] (36 and 72 Pa) and nitrogen supply (N; low and high) for 141 days in phytotron chambers. In oak, there was no significant effect of [CO₂] on wood structure, although elevated [CO₂] tended to decrease stomatal conductance (g _s) and increased water use efficiency regardless of the N treatment. However, high N supply increased root biomass and induced wider earlywood and larger vessels in the secondary xylem in stems, leading to increased hydraulic conductance. In alder, there was significant interactive effect of [CO₂] and N on vessel density, and high N supply increased the mean vessel area. Our results suggest that wood structures related to water transport were not markedly altered, although elevated [CO₂] induced changes in physiological parameters such as g _s and biomass allocation, and that N fertilization had more pronounced effects on non-N₂-fixing oak than on N₂-fixing alder.     

Production,decomposition, and nitrogen dynamics ofMyrica gale litter

Summary Myrica gale litter deposition and decomposition were studied in a central Massachusetts peatland to determine the amount of N made available to the ecosystem by these processes. Leaf litter added 114–140 g biomass m^–2 annually and contained 2.12–2.59 g N m^–2 returning about 70% as much N to the ecosystem as was fixed annually byMyrica gale. During the first five years of decomposition, the leaf liter lost only 40% of its initial biomass and released only 10% of its initial N content. About 60% of its original N mass was still present when the litter reached the permanently waterlogged zone, and thus was effectively lost to the vegetation. The low decomposition rate was due primarily to the chemical content of the litter because similarly low rates were observed in an upland forest where the native litter decayed rapidly. The initial lignin content (40%) ofM. gale litter may be largely responsible for its slow decomposition in spite of its relatively high (1.69%) initial N content.M. gale litter decayed substantially more slowly and had a much higher initial lignin content than the litter of other woody N₂-fixing plants which have been examined.