Towards an ecological understanding of biological nitrogen fixation

Biogeochemistry - N limitation to primary production and other ecosystem processes is widespread. To understand the causes and distribution of N limitation, we must understand the controls of...     

Nutrient Limitation to Nitrogen Fixation in Young Volcanic Sites

I used measures of ¹⁵N natural abundance and of nitrogenase activity (acetylene reduction) to examine whether the supply of non-N nutrients limits rates of N₂ fixation on young volcanic substrates in Hawaii. Leaves of the dominant tree (Metrosideros polymorpha, a nonfixer) were strongly depleted in ¹⁵N in control plots (–10.8 to –11.1⁰/₀₀). More than 5 y of repeated fertilization with P increased δ¹⁵N to –8.9 to –9.9⁰/₀₀, and the addition of all other essential plant nutrients (except N) together with P further increased ¹⁵N to –8.1 to –9.3⁰/₀₀. This pattern is consistent with enhanced N₂ fixation, because newly fixed N would have a δ¹⁵N near 0⁰/₀₀. Assays of nitrogenase activity in the experimental plots demonstrated that potential N fixation associated with nonvascular plants and with tree and fern litter were increased significantly by additions of P and by the combined nutrient treatment; when these were added together, the increase in nitrogenase activity was 6- to 11-fold over control plots. The supply of P and other weathering-derived nutrients constrains rates of N₂ fixation in these young volcanic sites and thereby contributes to the maintenance of N limitation to primary production and other ecosystem processes. Received 7 January 1999; accepted 3 May 1999.     

Ecosystem constraints to symbiotic nitrogen fixers: a simple model and its implications

The widespread occurrence of N limitation to net primary production (NPP) and other ecosystem processes, despite the ubiquitous occurrence of N-fixing symbioses, remains a significant puzzle in terrestrial ecology. We describe a simple simulation model for an ecosystem containing a generic nonfixer and a symbiotic N fixer, based on: (1) a higher cost for N acquisition by N fixers than nonfixers; (2) growth of fixers and fixation of N only when low N availability limits the growth of nonfixers, and other resources are available; and (3) losses of fixed N from the system only when the quantity of available N exceeds plant and microbial demands. Despite the disadvantages faced by the N fixer under these conditions, N fixation and loss adjust N availability close to the availability of other resources, and biomass and NPP in this simple model can be substantially but only transiently N limited. We then modify the model by adding: (1) losses of N in forms other than excess available N (e.g., dissolved organic N, trace gases produced by nitrification); and (2) constraints to the growth and activity of N fixers imposed by differential effects of shading, P limitation, and grazing. The combination of these processes is sufficient to describe an open system, with input from both precipitation and N fixation, that is nevertheless strongly N-limited at equilibrium. This model is useful for exploring causes and consequences of constraints to N fixation, and hence of N limitation, and we believe it will also be useful for evaluating how N fixation and limitation interact with elevated CO₂ and other components of global enviromental change.     

Anion fluxes in three Indiana forests

Summary The major ions in soil solution in control and disturbed plots were measured over a year in three Indiana forests. The major ions in bulk precipitation were hydrogen and sulfate. Sulfate, calcium, and magnesium were the most important ions in the soil solution in control plots, while in disturbed plots nitrate, calcium, and magnesium were most important. Increase nitrate production and loss following disturbance caused a large increase in cation losses.     

Potential ecosystem-level effects of genetic variation among populations of Metrosideros polymorpha from a soil fertility gradient in Hawaii

This study assessed intrinsic differences in tissue quality and growth rate among populations of Metrosideros polymorpha native to sites with a range of soil fertilities. We collected seedlings from three Hawaiian mesic forests that were either phosphorus-limited, nitrogen-limited, or relatively fertile. These individuals were grown in a common garden under a factorial high/low, N/P fertilization regime for 1.5 years and then harvested to determine genetic divergence; aboveground growth rate; and lignin, N, and P concentrations in leaves and roots. Allozyme analyses indicated that the three groups had genetically diverged to some degree (genetic distance = 0.036-0.053 among populations). Relative growth rate did not differ significantly among the populations. Senescent leaves from the fertile-site population had the highest N concentrations (due to low N resorption) and had lower lignin concentrations than plants from the N-limited site. Across treatments, P concentrations in senescent leaves were highest in plants from the fertile and P-limited site. Root tissue quality did not generally differ significantly among populations. Since decomposition rate of senescent leaves in this system is related positively to N concentration and negatively to lignin concentration, senescent leaves from the fertile-site population may have a genetic tendency toward faster decay than the others. The intrinsic qualities of the three populations may provide positive feedbacks on nutrient cycling at each site-nutrient availability may be raised to some degree at the fertile site, and reduced at the N- or P-limited sites. Our results suggest that even a small degree of genetic differentiation among groups can influence traits related to nutrient cycling.     

Vegetation Effects on Soil Organic Matter Chemistry of Aggregate Fractions in a Hawaiian Forest

We examined chemical changes from leaf tissue to soil organic matter (SOM) to determine the persistence of plant chemistry into soil aggregate fractions. We characterized a slow (Dicranopteris linearis) and fast-decomposing species (Cheirodendron trigynum) and surface (O), and subsurface (A-horizon) SOM beneath each species using pyrolysis-gas chromatography/mass spectrometry (py-GC/MS), with and without derivatization. The live tissues of Dicranopteris had greater lignin content whereas Cheirodendron had a greater lipid, N-bearing, and polysaccharide component. Despite this difference in leaf chemistry, SOM chemistry was similar between soil aggregate fractions, but different between horizons. The O-horizon contained primarily lignin and polysaccharide biomarkers whereas the A-horizon contained polysaccharide, aromatic, and N-derived compounds, indicating considerable microbial processing of plant litter. The soils beneath Cheirodendron inherited a greater lipid signal composed of cutin and suberin biomarkers whereas the soils beneath Dicranopteris contained greater aromatic biomarker content, possibly derived from plant lignins. The soils beneath both species were more similar to root polysaccharides, lipids, and lignins than aboveground tissue. This study indicates that although plant-derived OM is processed vigorously, species-specific biomarkers and compound class differences persist into these soils and that differences in plant chemical properties may influence soil development even after considerable reworking of plant litter by microorganisms.     

Ecosystem development on Hawaiian lava flows: biomass and species composition

Abstract. The strong environmental gradients and ‘natural experimental design’ of Mauna Loa volcano, Hawaii, provide an outstanding opportunity to study controls on ecosystem development. We measured above-ground vascular plant biomass and species composition on 42 sites on which precipitation, temperature, substrate texture, and substrate age varied substantially and largely independently. Biomass and species richness of live plants were strongly correlated with precipitation and lava flow age, but not with temperature or lava flow texture. Species composition, as measured by correspondence analysis, was likewise correlated with precipitation and flow age, but composition was also strongly influenced by temperature. Lava texture had a complex effect on vegetation, with ‘a’ a lava favoring vegetation development on wet sites and pāhoehoe favoring development on dry sites. Many locations remain virtually free of invasion by alien species; aliens appear where disturbance has facilitated invasion, either from stand-level dieback in rainforest or a grass-fire cycle on the dry, leeward side of the mountain. All four of the environmental factors studied here (precipitation, temperature, substrate texture, and substrate age) exert significant and independent control over vegetation biomass and/or species composition on Mauna Loa.