Nina Wurzburger

Potassium, phosphorus, or nitrogen limit root allocation, tree growth, or litter production in a lowland tropical forest

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 x 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

Hemlock Declines Rapidly with Hemlock Woolly Adelgid Infestation: Impacts on the Carbon Cycle of Southern Appalachian Forests

The recent infestation of southern Appalachian eastern hemlock stands by hemlock woolly adelgid (HWA) is expected to have dramatic and lasting effects on forest structure and function. We studied the short-term changes to the carbon cycle in a mixed stand of hemlock and hardwoods, where hemlock was declining due to either girdling or HWA infestation. We expected that hemlock would decline more rapidly from girdling than from HWA infestation. Unexpectedly, in response to both girdling and HWA infestation, hemlock basal area increment (BAI) reduced substantially compared to reference hardwoods in 3xa0years. This decline was concurrent with moderate increases in the BAI of co-occurring hardwoods. Although the girdling treatment resulted in an initial pulse of hemlock needle inputs, cumulative litter inputs and O horizon mass did not differ between treatments over the study period. Following girdling and HWA infestation, very fine root biomass declined by 20–40% in 2xa0years, which suggests hemlock root mortality in the girdling treatment, and a reduction in hemlock root production in the HWA treatment. Soil CO2 efflux (Esoil) declined by approximately 20% in 1xa0year after both girdling and HWA infestation, even after accounting for the intra-annual variability of soil temperature and moisture. The reduction in Esoil and the concurrent declines in BAI and standing very fine root biomass suggest rapid declines in hemlock productivity from HWA infestation. The accelerated inputs of detritus resulting from hemlock mortality are likely to influence carbon and nutrient fluxes, and dictate future patterns of species regeneration in these forest ecosystems.

Molybdenum and phosphorus interact to constrain asymbiotic nitrogen fixation in tropical forests.

Biological di-nitrogen fixation (N2) is the dominant natural source of new nitrogen to land ecosystems. Phosphorus (P) is thought to limit N2 fixation in many tropical soils, yet both molybdenum (Mo) and P are crucial for the nitrogenase reaction (which catalyzes N2 conversion to ammonia) and cell growth. We have limited understanding of how and when fixation is constrained by these nutrients in nature. Here we show in tropical forests of lowland Panama that the limiting element on asymbiotic N2 fixation shifts along a broad landscape gradient in soil P, where Mo limits fixation in P-rich soils while Mo and P co-limit in P-poor soils. In no circumstance did P alone limit fixation. We provide and experimentally test a mechanism that explains how Mo and P can interact to constrain asymbiotic N2 fixation. Fixation is uniformly favored in surface organic soil horizons - a niche characterized by exceedingly low levels of available Mo relative to P. We show that soil organic matter acts to reduce molybdate over phosphate bioavailability, which, in turn, promotes Mo limitation in sites where P is sufficient. Our findings show that asymbiotic N2 fixation is constrained by the relative availability and dynamics of Mo and P in soils. This conceptual framework can explain shifts in limitation status across broad landscape gradients in soil fertility and implies that fixation depends on Mo and P in ways that are more complex than previously thought.

Fine-root responses to fertilization reveal multiple nutrient limitation in a lowland tropical forest

Questions remain as to which soil nutrients limit primary production in tropical forests. Phosphorus (P) has long been considered the primary limiting element in lowland forests, but recent evidence demonstrates substantial heterogeneity in response to nutrient addition, highlighting a need to understand and diagnose nutrient limitation across diverse forests. Fine-root characteristics including their abundance, functional traits, and mycorrhizal symbionts can be highly responsive to changes in soil nutrients and may help to diagnose nutrient limitation. Here, we document the response of fine roots to long-term nitrogen (N), P, and potassium (K) fertilization in a lowland forest in Panama. Because this experiment has demonstrated that N and K together limit tree growth and P limits fine litter production, we hypothesized that fine roots would also respond to nutrient addition. Specifically we hypothesized that N, P, and K addition would reduce the biomass, diameter, tissue density, and mycorrhizal colonization of fine roots, and increase nutrient concentration in root tissue. Most morphological root traits responded to the single addition of K and the paired addition of N and P, with the greatest response to all three nutrients combined. The addition of N, P, and K together reduced fine-root biomass, length, and tissue density, and increased specific root length, whereas root diameter remained unchanged. Nitrogen addition did not alter root N concentration, but P and K addition increased root P and K concentration, respectively. Mycorrhizal colonization of fine roots declined with N, increased with P, and was unresponsive to K addition. Although plant species composition remains unchanged after 14 years of fertilization, fine-root characteristics responded to N, P, and K addition, providing some of the strongest stand-level responses in this experiment. Multiple soil nutrients regulate fine-root abundance, morphological and chemical traits, and their association with mycorrhizal fungi in a species-rich lowland tropical forest.

Nitrogen and phosphorus interact to control tropical symbiotic N2 fixation: a test in Inga punctata

Summary n nxa0Symbiotic di-nitrogen (N2) fixation is a critical biogeochemical process in tropical forests, yet it remains unresolved how fixation is controlled by the availability of soil nitrogen and phosphorus, two nutrients often considered limiting in terrestrial ecosystems. nxa0We examine whether individual N2-fixing trees can overcome nitrogen and phosphorus constraints by employing different strategies of nutrient acquisition and use: N2 fixation, phosphatase exudation, mycorrhizal symbiosis and changes in root–shoot ratio or tissue stoichiometry. nxa0We grew a common and widespread N2 fixer, Inga punctata, in a full factorial nitrogen and phosphorus addition experiment (each nutrient at three levels) and evaluated whether trees adjusted their strategies of nutrient acquisition to overcome limitation. nxa0N2 fixation was controlled by nitrogen availability in phosphorus-sufficient soils, but both fixation and plant growth were constrained by phosphorus in the unamended native phosphorus-poor soils. Despite the investment in both extracellular phosphatases and mycorrhizal symbionts, plants were unable to overcome phosphorus limitation. nxa0Our findings support the hypotheses that: (i) N2 fixation is proximately controlled by nitrogen availability, consistent with a facultative fixation strategy, and (ii) N2 fixation and N2 fixer biomass growth are ultimately constrained by soil phosphorus. We found no support for the hypothesis that fixers can overcome phosphorus limitation by trading fixed N2 for soil phosphorus. nxa0Synthesis. This study provides new knowledge about how nitrogen and phosphorus interact to regulate tropical N2 fixation by examining a suite of strategies that plants may employ to overcome nutrient limitation. These findings, focused at the organismal level, have broader implications for biogeochemical controls at the ecosystem level in tropical forests.

Drought enhances symbiotic dinitrogen fixation and competitive ability of a temperate forest tree

AbstractnGeneral circulation models project more intense and frequent droughts over the next century, but many questions remain about how terrestrial ecosystems will respond. Of particular importance, is to understand how drought will alter the species composition of regenerating temperate forests wherein symbiotic dinitrogen (N2)-fixing plants play a critical role. In experimental mesocosms we manipulated soil moisture to study the effect of drought on the physiology, growth and competitive interactions of four co-occurring North American tree species, one of which (Robinia pseudoacacia) is a symbiotic N2-fixer. We hypothesized that drought would reduce growth by decreasing stomatal conductance, hydraulic conductance and increasing the water use efficiency of species with larger diameter xylem vessel elements (Quercus rubra, R. pseudoacacia) relative to those with smaller elements (Acer rubrum and Liriodendron tulipifera). We further hypothesized that N2 fixation by R. pseudoacacia would decline with drought, reducing its competitive ability. Under drought, growth declined across all species; but, growth and physiological responses did not correspond to species’ hydraulic architecture. Drought triggered an 80xa0% increase in nodule biomass and N accrual for R. pseudoacacia, improving its growth relative to other species. These results suggest that drought intensified soil N deficiency and that R. pseudoacacia’s ability to fix N2 facilitated competition with non-fixing species when both water and N were limiting. Under scenarios of moderate drought, N2 fixation may alleviate the N constraints resulting from low soil moisture and improve competitive ability of N2-fixing species, and as a result, supply more new N to the ecosystem.

Ericoid mycorrhizal root fungi and their multicopper oxidases from a temperate forest shrub

Ericoid mycorrhizal fungi (ERM) may specialize in capturing nutrients from their hosts litter as a strategy for regulating nutrient cycles in terrestrial ecosystems. In spite of their potential significance, we know little about the structure of ERM fungal communities and the genetic basis of their saprotrophic traits (e.g., genes encoding extracellular enzymes). Rhododendron maximum is a model ERM understory shrub that influences the nutrient cycles of montane hardwood forests in the southern Appalachians (North Carolina, USA). We sampled ERM roots of R. maximum from organic and mineral soil horizons and identified root fungi by amplifying and sequencing internal transcribed spacer (ITS) ribosomal DNA (rDNA) collected from cultures and clones. We observed 71 fungal taxa on ERM roots, including known symbionts Rhizoscyphus ericae and Oidiodendron maius, putative symbionts from the Helotiales, Chaetothyriales, and Sebacinales, ectomycorrhizal symbionts, and saprotrophs. Supporting the idea that ERM fungi are adept saprotrophs, richness of root-fungi was greater in organic than in mineral soil horizons. To study the genetic diversity of oxidative enzymes that contribute to decomposition, we amplified and sequenced a portion of genes encoding multicopper oxidases (MCOs) from ERM ascomycetes. Most fungi possessed multiple copies of MCO sequences with strong similarities to known ferroxidases and laccases. Our findings indicate that R. maximum associates with a taxonomically and ecologically diverse fungal community. The study of MCO gene diversity and expression may be useful for understanding how ERM root fungi regulate the cycling of nutrients between the host plant and the soil environment.

Ectomycorrhizal fungal community structure across a bog-forest ecotone in southeastern Alaska

We examined the ectomycorrhizal (ECM) fungal community across a bog-forest ecotone in southeastern Alaska. The bog and edge were both characterized by poorly drained Histosols and a continuous layer of Sphagnum species, ericaceous shrubs, Carex species, and shore pine [Pinus contorta Dougl. ex Loud. var. contorta]. The forest had better-drained Inceptisols and Spodosols, a tree community comprised of western hemlock [Tsuga heterophylla (Raf.) Sarg.], yellow cedar (Thuja plicata Donn ex D. Don.), Sitka spruce [Picea sitchensis (Bong.) Carr.] and shore pine, and an understorey of ericaceous shrubs and herbs. ECM root tip density (tips cm−3 soil) was significantly greater in the forest than the edge or bog and ECM colonization was significantly different in all three plant communities. The below ground ECM fungal taxa were analyzed using molecular techniques (PCR-RFLP and DNA sequencing). Three ECM fungal taxa, Suillus tomentosus (Kauffman) Singer, Cenococcum geophilum Fr.:Fr, and a Russula species, differed in relative frequency, yet were among the four most frequent in all three plant communities. Although differences in ECM fungal richness were observed across plant communities, unequal sampling of ECM roots due to root density and colonization differences confounded richness comparisons. Using resampling procedures for creating taxon-accumulation curves as a function of sampled ECM roots revealed similarities in cumulative ECM fungal taxa richness across the ecotone.

Mycorrhizal associations of trees have different indirect effects on organic matter decomposition

Summary n1.Organic matter decomposition is the main process by which carbon (C) is lost from terrestrial ecosystems, and mycorrhizal associations of plants (i.e., arbuscular mycorrhizae (AM) and ectomycorrhizae (ECM)) may have different indirect effects on this loss pathway. AM and ECM plants differ in the soil decomposers they promote and the quality of litter they produce, which may result in different patterns of organic matter decomposition, and hence, soil C loss. n n2.To determine how mycorrhizal associations indirectly affect decomposer activity, we collected soils and litters from four AM and four ECM tree species from a mixed-deciduous temperate forest for a field and lab study. We first characterized in situ patterns in soil chemistry and soil microbial biomass among these eight tree species. We then conducted a microcosm experiment with mineral soils, leaf-litter and fine roots originating from these tree species, where we reciprocally crossed litters and soils, and quantified the rate of heterotrophic respiration over a 140 day lab incubation. n n3.In natural forest conditions, AM tree soils contained lower total C and microbial biomass C:N relative to ECM tree soils. In our microcosm experiment, AM soils supported greater heterotrophic respiration than did ECM soils. The addition of AM litter stimulated respiration more than did ECM litter, owing to the lower C:N of AM litter. Matching the mycorrhizal identity of litter and soil resulted in a difference in total respiration, such that combinations of AM litters with AM soils lost more C than did combinations of ECM litters and ECM soils. n n4.Synthesis Our findings demonstrate that AM and ECM trees have differing indirect effects on decomposer activity through the decomposers they cultivate and/or the quality of organic matter they produce in soils. Mycorrhizal differences in litter quality accentuate these effects on soil C loss and may explain patterns in soil C dynamics in terrestrial ecosystems. n nThis article is protected by copyright. All rights reserved.

Taxonomic identity determines N2 fixation by canopy trees across lowland tropical forests

Legumes capable of fixing atmospheric N2 are abundant and diverse in many tropical forests, but the factors determining ecological patterns in fixation are unresolved. A long-standing idea is that fixation depends on soil nutrients (N, P or Mo), but recent evidence shows that fixation may also differ among N2-fixing species. We sampled canopy-height trees across five species and one species group of N2-fixers along a landscape P gradient, and manipulated P and Mo to seedlings in a shadehouse. Our results identify taxonomy as the major determinant of fixation, with P (and possibly Mo) only influencing fixation following tree-fall disturbances. While 44% of trees did not fix N2, other trees fixed at high rates, with two species functioning as superfixers across the landscape. Our results raise the possibility that fixation is determined by biodiversity, evolutionary history and species-specific traits (tree growth rate, canopy stature and response to disturbance) in the tropical biome.