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"Fahey, Timothy"
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Soil nitrogen affects phosphorus recycling: foliar resorption and plant-soil feedbacks in a northern hardwood forest
by
See, Craig R.
,
Vadeboncoeur, Matthew A.
,
Fahey, Timothy J.
in
Acer rubrum
,
Acer saccharum
,
Betula alleganiensis
2015
Previous studies have attempted to link foliar resorption of nitrogen and phosphorus to their respective availabilities in soil, with mixed results. Based on resource optimization theory, we hypothesized that the foliar resorption of one element could be driven by the availability of another element. We tested various measures of soil N and P as predictors of N and P resorption in six tree species in 18 plots across six stands at the Bartlett Experimental Forest, New Hampshire, USA. Phosphorus resorption efficiency (
P
< 0.01) and proficiency (
P
= 0.01) increased with soil N content to 30 cm depth, suggesting that trees conserve P based on the availability of soil N. Phosphorus resorption also increased with soil P content, which is difficult to explain based on single-element limitation, but follows from the correlation between soil N and soil P. The expected single-element relationships were evident only in the O horizon: P resorption was high where resin-available P was low in the Oe (
P
< 0.01 for efficiency,
P
< 0.001 for proficiency) and N resorption was high where potential N mineralization in the Oa was low (
P
< 0.01 for efficiency and 0.11 for proficiency). Since leaf litter is a principal source of N and P to the O horizon, low nutrient availability there could be a result rather than a cause of high resorption. The striking effect of soil N content on foliar P resorption is the first evidence of multiple-element control on nutrient resorption to be reported from an unmanipulated ecosystem.
Journal Article
Incorporation of fine root detritus into forest soil organic matter
2023
One of the principal inputs of organic matter to forest soils is turnover of tree fine roots, but the process of decomposition of fine root litter and its conversion into stable soil organic matter (SOM) has received limited study. We labeled fine roots of sugar maple (Acer saccharum Marsh.) with 13C and traced the label for 7 years into four contrasting soils to improve understanding of this process. After 7 years we recovered an average of 8.9% of the 13C label, with about two-thirds recovered as coarse particulate organic matter and one-third in microaggregates and on silt and clay particles. No differences in 13C recovery were detected between 1–2 and 3–4 order fine roots. Most of the 13C in microaggregates (53–250 µm, 58%) was occluded within macroaggregates, and the recovery in this fraction increased significantly from year 2 to 7, illustrating the role of fine root detritus in the formation of microaggregates. This process was most pronounced in the A horizon of a higher pH soil (pH = 5.5) with high iron oxide content. Conversely, the lowest 13C recovery in this fraction was observed in the A horizon of an acidic, fine-textured Inceptisol (Cambisol—World Reference Base). We estimate that annual input into relatively stable fractions of SOM represents about 14% of the total annual accumulation in these fractions; thus, our results support recent evidence that fine root litter is only a moderate contributor to stable SOM in acid temperate forest soils.
Journal Article
Co-limitation of Fine Root Growth by Nitrogen and Phosphorus in Early Successional Northern Hardwood Forests
2024
Functional balance theory predicts that plants will allocate less carbon belowground when the availability of nutrients is elevated. We tested this prediction in two successional northern hardwood forest stands by quantifying fine root biomass and growth after 5–7 years of treatment in a nitrogen (N) x phosphorus (P) factorial addition experiment. We quantified root responses at two different levels of treatment: the whole-plot scale fertilization and small-patch scale fertilization of ingrowth cores. Fine root biomass was higher in plots receiving P, and fine root growth was highest in plots receiving both N and P. Thus, belowground productivity did not decrease in response to long-term addition of nutrients. We did not find conclusive evidence that elevated availability of one nutrient at the plot scale induced foraging for the other nutrient at the core scale, or that foraging for nutrients at the core scale responded to addition of limiting nutrients. Our observations suggest NP co-limitation of fine root growth and indicate complex interactions of N and P affecting aboveground and belowground production in early successional northern hardwood forest ecosystems.
Journal Article
Experimental approach and initial forest response to a simulated ice storm experiment in a northern hardwood forest
by
Frank Bowles
,
Wendy Leuenberger
,
Gary J. Hawley
in
Accretion
,
Biology and Life Sciences
,
Boreal forests
2020
Ice storms are a type of extreme winter weather event common to north temperate and boreal forests worldwide. Recent climate modelling studies suggest that these storms may become more frequent and severe under a changing climate. Compared to other types of storm events, relatively little is known about the direct and indirect impacts of these storms on forests, as naturally occurring ice storms are inherently difficult to study. Here we describe a novel experimental approach used to create a suite of ice storms in a mature hardwood forest in New Hampshire, USA. The experiment included five ice storm intensities (0, 6.4, 12.7 and 19.1 mm radial ice accretion) applied in a single year, and one ice storm intensity (12.7 mm) applied in two consecutive years. Results demonstrate the feasibility of this approach for creating experimental ice storms, quantify the increase in fine and coarse woody debris mass and nutrients transferred from the forest canopy to the soil under the different icing conditions, and show an increase in the damage to the forest canopy with increasing icing that evolves over time. In this forest, little damage occurred below 6.4 mm radial ice accretion, moderate damage occurred with up to 12.7 mm of accretion, and significant branch breakage and canopy damage occurred with 19.1 mm of ice. The icing in consecutive years demonstrated an interactive effect of ice storm frequency and severity such that some branches damaged in the first year of icing appeared to remain in the canopy and then fall to the ground in the second year of icing. These results have implications for National Weather Service ice storm warning levels, as they provide a quantitative assessment of ice-load related inputs of forest debris that will be useful to municipalities creating response plans for current and future ice storms.
Journal Article
Fine Root Growth Increases in Response to Nitrogen Addition in Phosphorus-limited Northern Hardwood Forests
2022
Resource allocation theory posits that increased soil nutrient availability results in decreased plant investment in nutrient acquisition. We evaluated this theory by quantifying fine root biomass and growth in a long term, nitrogen (N) × phosphorus (P) fertilization study in three mature northern hardwood forest stands where aboveground growth increased primarily in response to P addition. We did not detect a decline in fine root biomass or growth in response to either N or P. Instead, fine root growth increased in response to N, by 40% for length (P = 0.04 for the main effect of N in ANOVA), and by 36% for mass, relative to controls. Fine root mass growth was lower in response to N + P addition than predicted from the main effects of N and P (P = 0.01 for the interaction of N × P). The response of root growth to N availability did not result in detectable responses in fine root biomass (P = 0.61), which is consistent with increased root turnover with N addition. We propose that the differential growth response to fertilization between above- and belowground components is a mechanism by which trees enhance P acquisition in response to increasing N availability, illustrating how both elements may co-limit northern hardwood forest production.
Journal Article
How Much Soil Carbon is Derived from Woody Detritus? A Ten-Year Study of 13C Incorporation into Soil Organic Matter
by
Mathisson, Rachel
,
Fahey, Timothy J
,
Heinz, Alexis K
in
Biodiesel fuels
,
Biofuels
,
Carbon dioxide
2024
The importance of woody detritus as a source of soil organic matter is not well constrained. We quantified the recovery of 13C derived from isotopic-enriched sugar maple wood in various C fractions of two temperate forest soils in central New York, USA. Decay rates of small woody debris were quite rapid (k = 0.362 to 0.477 per year) and after 10 years less than 1% of the original wood mass remained in incubation bags. After six years we recovered only 0.26% (± 0.025) of the added 13C in the upper 5 cm of underlying soil. After 10 years this recovery declined to 0.11% (± 0.020) indicating substantial lability of retained SOC; most of this decline occurred from year 6 to 8 in the 1–5 cm depth increment, suggesting that the residue was quite stable at 10 years. The largest fraction of 13C was recovered in microaggregates (45%), especially those occluded within macroaggregates (30%), with a smaller proportion associated with the silt + clay fraction (20%). These proportions did not change significantly from year 6 to 10. Faster decay and higher 13C recovery were coincident with abundant saproxylic invertebrates from Scarabaeidae at one of the sites. We conclude that small woody debris is a minor source of stable SOC in these temperate forests (that is, less than 1% of annual SOC accumulation).
Journal Article
Treatment effects of nitrogen and phosphorus addition on foliar traits in six northern hardwood tree species
by
Vadeboncoeur, Matthew A.
,
Fahey, Timothy J.
,
Yanai, Ruth D.
in
Acer
,
Acer rubrum
,
Acer saccharum
2025
Foliar traits can reflect fitness responses to environmental changes, such as changes in nutrient availability. Species may respond differently to these changes due to differences in traits and their plasticity. Traits and community composition together can influence forest nutrient cycling. We compared five traits—foliar N, foliar P, specific leaf area (SLA), leaf dry matter content (LDMC), and leaf carbon isotope ratio (δ
13
C)—in six northern hardwood tree species (
Acer rubrum
,
Acer saccharum
,
Betula alleghaniensis
,
Betula papyrifera
,
Fagus grandifolia
, and
Prunus pensylvanica
) in a nitrogen (N) and phosphorus (P) fertilization study across 10 mid- and late-successional forest stands in New Hampshire, USA. We also analyzed the response of tree growth to N and P addition. Nutrient addition shifted trait values towards the “acquisitive” side of the spectrum for all traits except δ
13
C, reflecting a tradeoff between water-use efficiency and nutrient-use efficiency. Treatment responses in relative basal area increment revealed that the
Betula
species were N-limited, but traits of all species responded to either or both N and P addition in ways that suggest N and P co-limitation. Two species displayed lower foliar P under N addition, and three species displayed lower foliar N under P addition, which also suggests co-limitation. These indications of co-limitation were reflected at the community level. Specific leaf area, LDMC, and δ
13
C differed with stand age within several species. Examining trait responses of tree species and communities to nutrient availability increases our understanding of biological mechanisms underlying the complex effects of nutrient availability on forests.
Journal Article
Root morphology and architecture respond to N addition in Pinus tabuliformis, west China
by
Fahey, Timothy J.
,
liu, Fang
,
Xue, Sha
in
Animal and plant ecology
,
Animal, plant and microbial ecology
,
Anthropogenic factors
2013
Belowground dynamics of terrestrial ecosystems are responding to global increases in anthropogenic N deposition with important consequences for productivity and ecosystem health. We compared root characteristics across five root orders in Pinus tabuliformis plantations treated for 3 years to a gradient of N addition (0—15 g m -2 year -1 ). In reference plots, the roots of P. tabuliformis were finer and with higher specific root length than reported for other pine species, suggesting severe N limitation. Addition of N resulted in slightly reduced fine root biomass and significant changes in root morphology, responses that were associated primarily with first and second order roots. In particular, root number, cumulative root length, individual root length, and specific root length all declined with increasing N addition for first and second order roots, with most of the responses elicited at <9 g m -2 year -1 N addition. These responses (1) support the concept of ephemeral root modules consisting of first and second orders and (2) are consistent with a change in functional demand from uptake to transport with increasing soil resource availability. Traditionally, fine roots have been identified by a somewhat arbitrary diameter cut-off (e.g., 1 or 2 mm); as an index of fine root function, diameter would fail to reveal most of the functional response.
Journal Article
Phosphorus limitation of aboveground production in northern hardwood forests
by
Garrison-Johnston, Mariann
,
Vadeboncoeur, Matthew A.
,
Fahey, Timothy J.
in
aboveground productivity
,
age structure
,
Anthropogenic factors
2018
Forest productivity on glacially derived soils with weatherable phosphorus (P) is expected to be limited by nitrogen (N), according to theories of long-term ecosystem development. However, recent studies and model simulations based on resource optimization theory indicate that productivity can be co-limited by N and P. We conducted a full factorial N × P fertilization experiment in 13 northern hardwood forest stands of three age classes in central New Hampshire, USA, to test the hypothesis that forest productivity is co-limited by N and P. We also asked whether the response of productivity to N and P addition differs among species and whether differential species responses contribute to community-level co-limitation. Plots in each stand were fertilized with 30 kg N·ha−1·yr−1, 10 kg P·ha−1·yr−1, N + P, or neither nutrient (control) for four growing seasons. The productivity response to treatments was assessed using per-tree annual relative basal area increment (RBAI) as an index of growth. RBAI responded significantly to P (P = 0.02) but not to N (P = 0.73). However, evidence for P limitation was not uniform among stands. RBAI responded to P fertilization in mid-age (P = 0.02) and mature (P = 0.07) stands, each taken as a group, but was greatest in N-fertilized plots of two stands in these age classes, and there was no significant effect of P in the young stands. Both white birch (Betula papyrifera Marsh.) and beech (Fagus grandifolia Ehrh.) responded significantly to P; no species responded significantly to N. We did not find evidence for N and P co-limitation of tree growth. The response to N + P did not differ from that to P alone, and there was no significant N × P interaction (P = 0.68). Our P limitation results support neither the N limitation prediction of ecosystem theory nor the N and P co-limitation prediction of resource optimization theory, but could be a consequence of long-term anthropogenic N deposition in these forests. Inconsistencies in response to P suggest that successional status and variation in site conditions influence patterns of nutrient limitation and recycling across the northern hardwood forest landscape.
Journal Article
Tree Species and Mycorrhizal Associations Influence the Magnitude of Rhizosphere Effects
2006
Previous research on the effects of tree species on soil processes has focused primarily on the role of leaf litter inputs. We quantified the extent to which arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) tree species influence soil microbial activity and nutrient availability through rhizosphere effects. Rhizosphere soil, bulk soil, and fine roots were collected from 12 monospecific plots (six AM and six ECM tree species) planted on a common soil at the Turkey Hill Plantations in Dryden, New York. Rhizosphere effects were estimated by the percentage difference between rhizosphere and bulk soil samples for several assays. Rhizosphere effects on soil microbes and their activities were significant for ECM species but in only a few cases for AM species. In AM tree species, microbial biomass, net N mineralization, and phosphatase enzyme activity in the rhizosphere were 10-12% greater than in bulk soil. In ECM tree species, rhizosphere effects for microbial biomass, C mineralization rates, net N mineralization, and phosphatase activity were 25-30% greater than bulk soil, and significantly greater than AM rhizosphere effects. The magnitude of rhizosphere effects was negatively correlated with the degree of mycorrhizal colonization in AM tree species (r = -0.83) and with fine root biomass (r = -0.88) in ECM tree species, suggesting that different factors influence rhizosphere effects in tree species forming different mycorrhizal associations. Rhizosphere effects on net N mineralization and phosphatase activity were also much greater in soils with pH < 4.3 for both AM and ECM tree species, suggesting that soil pH and its relation to nutrient availability may also influence the magnitude of rhizosphere effects. Our results support the idea that tree roots stimulate nutrient availability in the rhizosphere, and that systematic differences between AM and ECM may result in distinctive rhizosphere effects for C, N, and P cycling between AM and ECM tree species.
Journal Article