Nitrogen (N)‐fixing trees are thought to break a basic rule of leaf economics: higher leaf N concentrations do not translate into higher rates of carbon assimilation. Understanding how leaf N affects photosynthesis and water use efficiency (WUE) in this ecologically important group is critical. We grew six N‐fixing and four non‐fixing tree species for 4–5 years at four fertilization treatments in field experiments in temperate and tropical regions to assess how functional type (N fixer vs. non‐fixer) and N limitation affected leaf N and how leaf N affected light‐saturated photosynthesis (
Light and soil nitrogen availability can be strong controls of plant nitrogen (N) fixation, but data on how understory N‐fixing plants respond to these drivers are limited despite their important role in ecosystem N cycling. Furthermore, ecosystem N cycling can be altered by the introduction of species with nutrient use patterns that differ from natives. We assessed how N fixation of two exotic, understory species responded to varying light and soil N environments. We sampled leaf tissue from
- PAR ID:
- 10405368
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Journal of Ecology
- Volume:
- 111
- Issue:
- 4
- ISSN:
- 0022-0477
- Page Range / eLocation ID:
- p. 915-926
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract A sat), stomatal conductance (g sw) and WUE (WUEiand δ13C).A sat, WUEiand δ13C, but notg sw, increased with higher leaf N. Surprisingly, N‐fixing and non‐fixing trees displayed similar scaling between leaf N and these physiological variables, and this finding was supported by reanalysis of a global dataset. N fixers generally had higher leaf N than non‐fixers, even when non‐fixers were not N‐limited at the leaf level. Leaf‐level N limitation did not alter the relationship ofA sat,g sw, WUEiand δ13C with leaf N, although it did affect the photosynthetic N use efficiency. Higher WUE was associated with higher productivity, whereas higherA satwas not.Synthesis : The ecological success of N‐fixing trees depends on the effect of leaf N on carbon gain and water loss. Using a field fertilization experiment and reanalysis of a global dataset, we show that high leaf‐level photosynthesis and WUE in N fixers stems from their higher average leaf N, rather than a difference between N fixers and non‐fixers in the scaling of photosynthesis and WUE with leaf N. By clarifying the mechanism by which N fixers achieve and benefit from high WUE, our results further the understanding of global N fixer distributions. -
Abstract Symbiotic nitrogen fixation (SNF) is a key ecological process whose impact depends on the strategy of SNF regulation—the degree to which rates of SNF change in response to limitation by N versus other resources. SNF that is obligate or exhibits incomplete downregulation can result in excess N fixation, whereas a facultative SNF strategy does not. We hypothesized that tree‐based SNF strategies differed by latitude (tropical vs. temperate) and symbiotic type (actinorhizal vs. rhizobial). Specifically, we expected tropical rhizobial symbioses to display strongly facultative SNF as an explanation of their success in low‐latitude forests. In this study we used15N isotope dilution field experiments in New York, Oregon, and Hawaii to determine SNF strategies in six N‐fixing tree symbioses. Nitrogen fertilization with +10 and +15 g N m−2 year−1for 4–5 years alleviated N limitation in all taxa, paving the way to determine SNF strategies. Contrary to our hypothesis, all six of the symbioses we studied sustained SNF even at high N.
Robinia pseudoacacia (temperate rhizobial) fixed 91% of its N (%Ndfa) in controls, compared to 64% and 59% in the +10 and +15 g N m−2 year−1treatments. ForAlnus rubra (temperate actinorhizal), %Ndfawas 95%, 70%, and 60%. For the tropical species, %Ndfawas 86%, 80%, and 82% forGliricidia sepium (rhizobial); 79%, 69%, and 67% forCasuarina equisetifolia (actinorhizal); 91%, 42%, and 67% forAcacia koa (rhizobial); and 60%, 51%, and 19% forMorella faya (actinorhizal). Fertilization with phosphorus did not stimulate tree growth or SNF. These results suggest that the latitudinal abundance distribution of N‐fixing trees is not caused by a shift in SNF strategy. They also help explain the excess N in many forests where N fixers are common. -
Summary Nitrogen (N2)‐fixing moss microbial communities play key roles in nitrogen cycling of boreal forests. Forest type and leaf litter inputs regulate moss abundance, but how they control moss microbiomes and N2‐fixation remains understudied. We examined the impacts of forest type and broadleaf litter on microbial community composition and N2‐fixation rates of
Hylocomium splendens andPleurozium schreberi .We conducted a moss transplant and leaf litter manipulation experiment at three sites with paired paper birch (
Betula neoalaskana ) and black spruce (Picea mariana ) stands in Alaska. We characterized bacterial communities using marker gene sequencing, determined N2‐fixation rates using stable isotopes (15N2) and measured environmental covariates.Mosses native to and transplanted into spruce stands supported generally higher N2‐fixation and distinct microbial communities compared to similar treatments in birch stands. High leaf litter inputs shifted microbial community composition for both moss species and reduced N2‐fixation rates for
H. splendens , which had the highest rates. N2‐fixation was positively associated with several bacterial taxa, including cyanobacteria.The moss microbiome and environmental conditions controlled N2‐fixation at the stand and transplant scales. Predicted shifts from spruce‐ to deciduous‐dominated stands will interact with the relative abundances of mosses supporting different microbiomes and N2‐fixation rates, which could affect stand‐level N inputs.
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Abstract Carbon uptake by the terrestrial biosphere depends on supplies of new nitrogen (N) from symbiotic N fixation, but we lack a framework for scaling fixation accurately and for predicting its response to global change.
We scaled symbiotic N fixation from individual N fixers (i.e. plants that host N‐fixing bacteria), by quantifying three key parameters—the abundance of N fixers, whether they are fixing N and their N fixation rates. We apply this framework to black locust, a widespread N‐fixing tree in temperate forests of the eastern United States, and harness long‐term data from southern Appalachian forests to scale fixation from trees to the landscape and over succession.
Symbiotic N fixation at the landscape scale peaked in the first decade following forest disturbance, and then declined. This pattern was due to the declining density and declining fixation rates of individual black locust trees over succession. Independent of forest succession, and coincident with chronic atmospheric N deposition, we have evidence suggesting that nodule biomass produced by black locust trees has declined by 83% over the last three decades. This difference in nodule biomass translates to a maximum fixation rate of 11 kg N ha−1 year−1and a landscape average of 1.5 kg N ha−1 year−1in contemporary forests.
Synthesis . We find key controls on symbiotic N fixation by black locust over space and time, suggesting lower fixation rates in eastern deciduous forests than previous estimates. Our scaling framework can be applied to other N fixers to aid predictions of symbiotic N fixation and ecosystem response to global change. -
Abstract Most forests are recovering from human land use, making it critical to understand the effect of disturbance on forest recovery. Forests of the eastern United States have a long history of land use, but it is unknown whether historical disturbances have contributed to their transition from ectomycorrhizal (ECM) to arbuscular mycorrhizal (AM) tree dominance. Disturbance may promote nitrogen (N)‐fixing trees in early succession, which can elevate soil N availability even after they die. Higher soil N availability may facilitate the competitive success of AM trees over ECM trees, but such ‘N fixer founder effects’ have not been empirically tested.
Here, we analysed data from three land‐use disturbances in a temperate forest historically dominated by ECM trees: selective‐cutting (ranging from 0 to 52 m2 ha−1), clear‐cutting and agricultural abandonment. These disturbances occurred at different times, but long‐term data capture 3–7 decades of forest recovery.
We found that the AM tree fraction in contemporary forests was 2, 4, and 6‐fold higher following selective‐cutting, clear‐cutting and agricultural abandonment, respectively, compared to forest composition in 1934. Across these disturbances we also observed an increasing abundance of the N fixer black locust immediately following disturbance. Using a simulation model parameterized by data from black locust, we estimated historical rates of symbiotic N fixation to understand the relationship between N fixation and AM dominance in individual plots. We found that N fixation was positively associated with the growth of ECM trees generally, and oak and hickory specifically, only following light selective‐cutting (<12 or <18 m2 ha−1basal area extraction, respectively). Following higher levels of selective‐cutting and clear‐cutting, N fixation was positively associated with the growth of AM trees, particularly red maple and tulip poplar. Agricultural abandonment led to AM dominance regardless of N fixation rates.
Synthesis and applications . Our findings suggest that common land use practices and black locust, a native N fixer, can reduce the dominance of ECM trees. If N fixers are likely to proliferate following disturbance, we might maintain ECM dominance by cutting trees at low densities and by applying prescribed fire to remove N.