Allometric relationships for eight species of 4-5 year old nitrogen-fixing and non-fixing trees.

Allometric equations are often used to estimate plant biomass allocation to different tissue types from easier-to-measure quantities. Biomass allocation, and thus allometric equations, often differs by species and sometimes varies with nutrient availability. We measured biomass components for five nitrogen-fixing tree species (Robinia pseudoacacia, Gliricidia sepium, Casuarina equisetifolia, Acacia koa, Morella faya) and three non-fixing tree species (Betula nigra, Psidium cattleianum, Dodonaea viscosa) grown in field sites in New York and Hawaii for 4-5 years and subjected to four fertilization treatments. We measured total aboveground, foliar, main stem, secondary stem, and twig biomass in all species, and belowground biomass in Robinia pseudoacacia and Betula nigra, along with basal diameter, height, and canopy dimensions. The individuals spanned a wide size range (<1-16 cm basal diameter; 0.24-8.8 m height). For each biomass component, aboveground biomass, belowground biomass, and total biomass, we determined the following four allometric equations: the most parsimonious (lowest AIC) overall, the most parsimonious without a fertilization effect, the most parsimonious without canopy dimensions, and an equation with basal diameter only. For some species, the most parsimonious overall equation included fertilization effects, but fertilization effects were inconsistent across fertilization treatments. We therefore concluded that fertilization does not clearly affect allometric relationships in these species, size classes, and growth conditions. Our best-fit allometric equations without fertilization effects had the following R2 values: 0.91-0.99 for aboveground biomass (the range is across species), 0.95 for belowground biomass, 0.80-0.96 for foliar biomass, 0.94-0.99 for main stem biomass, 0.77-0.98 for secondary stem biomass, and 0.88-0.99 for twig biomass. Our equations can be used to estimate overall biomass and biomass of tissue components for these size classes in these species, and our results indicate that soil fertility does not need to be considered when using allometric relationships for these size classes in these species.

Nitrogen fixation in the stag beetle, Ceruchus piceus (Coleoptera: Lucanidae): could insects contribute more to ecosystem nitrogen budgets than previously thought?

Nitrogen (N) is a key nutrient required by all living organisms for growth and development, but is a limiting resource for many organisms. Organisms that feed on material with low N content, such as wood, might be particularly prone to N limitation. In this study, we investigated the degree to which the xylophagous larvae of the stag beetle Ceruchus piceus (Weber) use associations with N-fixing bacteria to acquire N. We paired acetylene reduction assays by cavity ring-down absorption spectroscopy (ARACAS) with 15N2 incubations to characterize rates of N fixation within C. piceus. Not only did we detect significant N fixation activity within C. piceus larvae, but we calculated a rate that was substantially higher than most previous reports for N fixation in insects. While taking these measurements, we discovered that N fixation within C. piceus can decline rapidly in a lab setting. Consequently, our results demonstrate that previous studies, which commonly keep insects in the lab for long periods of time prior to and during measurement, may have systematically under-reported rates of N fixation in insects. This suggests that within-insect N fixation may contribute more to insect nutrition and ecosystem-scale N budgets than previously thought.

Water balance affects foliar and soil nutrients differently.

Water balance influences soil development, and consequently plant communities, by driving weathering of soil minerals and leaching of plant nutrients from the soil. Along gradients in water balance, soils exhibit process domains where chemical properties are relatively stable punctuated by pedogenic thresholds where soil chemical properties change rapidly with little additional change in water balance. We ask if plant macronutrient concentrations in leaves also exhibit non-linear trends along water balance gradients, and if so, how these non-linearities relate to those in soils. We analyze foliar nutrient concentrations and foliar N:P ratios from eight species that span a range of growth forms along three water balance gradients (three of the species are found on multiple gradients). The gradients are located on basaltic substrate of different ages and have previously been characterized by studies on soil development. We find that maximum concentrations of foliar macronutrients occur at an intermediate water balance. As with soil nutrients, time mediates the effect of water balance on foliar nutrients, such that plants on older soils attain maximum nutrient concentrations at a lower water balance. On both a young, 20 ky and an old, 4100 ky water balance gradient, foliar nutrients reach peak concentrations at a water balance greater than the threshold for depletion of rock-derived nutrients in surface soils. Our findings suggest that plant acquisition of essential nutrients is imperfectly predicted by overall soil nutrient availability because the regulation of internal nutrient pools by plants makes nutrient pools within leaves partially independent of soil nutrient availability.

Temperature sensitivity of woody nitrogen fixation across species and growing temperatures.

The future of the land carbon sink depends on the availability of nitrogen (N)1,2 and, specifically, on symbiotic N fixation3-8, which can rapidly alleviate N limitation. The temperature response of symbiotic N fixation has been hypothesized to explain the global distribution of N-fixing trees9,10 and is a key part of some terrestrial biosphere models (TBMs)3,7,8, yet there are few data to constrain the temperature response of symbiotic N fixation. Here we show that optimal temperatures for N fixation in four tree symbioses are in the range 29.0-36.9 °C, well above the 25.2 °C optimum currently used by TBMs. The shape of the response to temperature is also markedly different to the function used by TBMs (asymmetric rather than symmetric). We also show that N fixation acclimates to growing temperature (hence its range of optimal temperatures), particularly in our two tropical symbioses. Surprisingly, optimal temperatures were 5.2 °C higher for N fixation than for photosynthesis, suggesting that plant carbon and N gain are decoupled with respect to temperature. These findings may help explain why N-fixing tree abundance is highest where annual maximum temperatures are >35 °C (ref. 10) and why N-fixing symbioses evolved during a warm period in the Earth's history11,12. Everything else being equal, our findings indicate that climate warming will probably increase N fixation, even in tropical ecosystems, in direct contrast to past projections8.