Stephen C. Hart

Biogeochemical Hot Spots and Hot Moments at the Interface of Terrestrial and Aquatic Ecosystems

Rates and reactions of biogeochemical processes vary in space and time to produce both hot spots and hot moments of elemental cycling. We define biogeochemical hot spots as patches that show disproportionately high reaction rates relative to the surrounding matrix, whereas hot moments are defined as short periods of time that exhibit disproportionately high reaction rates relative to longer intervening time periods. As has been appreciated by ecologists for decades, hot spot and hot moment activity is often enhanced at terrestrial-aquatic interfaces. Using examples from the carbon (C) and nitrogen (N) cycles, we show that hot spots occur where hydrological flowpaths converge with substrates or other flowpaths containing complementary or missing reactants. Hot moments occur when episodic hydrological flowpaths reactivate and/or mobilize accumulated reactants. By focusing on the delivery of specific missing reactants via hydrologic flowpaths, we can forge a better mechanistic understanding of the factors that create hot spots and hot moments. Such a mechanistic understanding is necessary so that biogeochemical hot spots can be identified at broader spatiotemporal scales and factored into quantitative models. We specifically recommend that resource managers incorporate both natural and artificially created biogeochemical hot spots into their plans for water quality management. Finally, we emphasize the needs for further research to assess the potential importance of hot spot and hot moment phenomena in the cycling of different bioactive elements, improve our ability to predict their occurrence, assess their importance in landscape biogeochemistry, and evaluate their utility as tools for resource management.

Competition for nitrogen between plants and soil microorganisms.

Experiments suggest that plants and soil microorganisms are both limited by inorganic nitrogen, even on relatively fertile sites. Consequently, plants and soil microorganisms may compete for nitrogen. While past research has focused on competition for inorganic nitrogen, recent studies have found that plants/mycorrhizae in a wide range of ecosystems can use organic nitrogen. A new view of competitive interactions between plants and soil microorganisms is necessary in ecosystem where plant uptake of organic nitrogen is observed.

Hydraulic lift: a potentially important ecosystem process

Hydraulic lift is the process by which some deep-rooted plants take in water from lower soil layers and exude that water into upper, drier soil layers. Hydraulic lift is beneficial to the plant transporting the water, and may be an important water source for neighboring plants. Recent evidence shows that hydraulically lifted water can promote greater plant growth, and could have important implications for net primary productivity, as well as ecosystem nutrient cycling and water balance.

INSECT HERBIVORY INCREASES LITTER QUALITY AND DECOMPOSITION: AN EXTENSION OF THE ACCELERATION HYPOTHESIS

Herbivore alteration of litter inputs may change litter decomposition rates and influence ecosystem nutrient cycling. In a semiarid woodland at Sunset Crater National Monument, Arizona, long-term insect herbivore removal experiments and the presence of herbivore resistant and susceptible pinyon pines (Pinus edulis) have allowed characterization of the population- and community-level effects of herbivory. Here we report how these same two herbivores, the mesophyll-feeding scale insect Matsucoccus acalyptus and the stem-boring moth Dioryctria albovittella alter litter quality, dynamics, and decomposition in this ecosystem. We measured aboveground litterfall, litter chemical composition, and first-year litter decomposition rates for trees resistant and susceptible to both herbivores and for susceptible trees from which herbivores had been experimentally removed for 16–18 years. Both herbivores significantly increased nitrogen concentration and decreased lignin:nitrogen and carbon:nitrogen ratios of abovegrou...

Physiological response to groundwater depth varies among species and with river flow regulation

We investigated the physiological response of two native riparian tree spe- cies (Populus fremontii and Salix gooddingii) and one exotic species (Tamarix chinensis) to groundwater availability along gradients of depth to groundwater at two rivers in Arizona. Depth to groundwater (DGW) at the dam-regulated Bill Williams River (BWR) was relatively constant and shallow (,4 m). Populus fremontiiat BWR did not experience reduced water availability at deeper groundwater depths, as evidenced by high predawn water potential. However, leaf gas exchange of P. fremontii was sensitive to high vapor pressure deficit where surface flow was ephemeral at BWR. Lower predawn water po- tentials of S. gooddingii at BWR suggested reduced water availability at deeper ground- water depths, but these reductions did not adversely affect net photosynthetic rate. Along the range of depth to groundwater at BWR, all three species suffered little canopy dieback, and dieback was not related to depth to groundwater. Depth to groundwater at the free- flowing Hassayampa River (HRP) was much greater and declined more rapidly in the ephemeral reaches than at BWR. Both P. fremontii and S. gooddingii experienced reduced water availability at deeper groundwater depths at HRP, as evidenced by lower predawn water potential. Both species also experienced reduced leaf gas exchange at deeper groundwater depths. Canopy dieback of all species was higher at HRP than at BWR and increased with increasing DGW, especially when DGW fell below 3 m. There was evidence to support branch sacrifice in these three riparian tree species as a means of improving water status in the surviving shoot. However, branch sacrifice was insufficient to prevent mortality in some of the native trees where DGW fell below 3 m at HRP. In contrast to the native species, T. chinensis showed no change in water availability, leaf gas exchange, or canopy dieback with increasing DGW at either river. Leaf gas exchange was lower and dieback was greater for T. chinensis at HRP where depth to groundwater was greater than at BWR, but there was no mortality at either river. Our results show that deep groundwater is more detrimental to the physiological condition of P. fremontii and S. gooddingii than it is to T. chinensis. Also, the pronounced differences in DGW and tree physiological performance between BWR and HRP suggest that dam regulation can in- crease water availability to mature trees in some desert riparian ecosystems. Finally, our study also provides estimates of the range of DGW that can maintain healthy, mature P. fremontii and S. gooddingii trees.

ECOLOGICAL RESTORATION ALTERS NITROGEN TRANSFORMATIONS IN A PONDEROSA PINE-BUNCHGRASS ECOSYSTEM

Ponderosa pine-bunchgrass ecosystems of the western United States were altered following Euro-American settlement as grazing and fire suppression facilitated pine invasion of grassy openings. Pine invasion changed stand structure and fire regimes, mo- tivating restoration through forest thinning and prescribed burning. To determine effects of restoration on soil nitrogen (N) transformations, we replicated (0.25-ha plots) the fol- lowing experimental restoration treatments within a ponderosa pine-bunchgrass community near Flagstaff, Arizona: (1) partial restoration—thinning to presettlement conditions, (2) complete restoration—removal of trees and forest floor to presettlement conditions, native grass litter addition, and a prescribed burn, and (3) control. Within treatments, we stratified sampling to assess effects of canopy cover on N transformations. Forest floor net N min- eralization and nitrification were similar among treatments on an areal basis, but higher in restoration treatments on a mass basis. In the mineral soil (0-15 cm), restoration treatments had 2-3 times greater annual net N mineralization and 3-5 times greater annual net nitri- fication than the control. Gross N transformation measurements indicate that elevated net N mineralization may be due to increased gross N mineralization, while elevated net ni- trification may be due to decreased microbial immobilization of nitrate. Net N transfor- mation rates beneath relict grassy openings were twice those beneath postsettlement pines. These short-term (1 yr) results suggest that ecological restoration increases N transformation rates and that prescribed burning may not be necessary to restore N cycling processes. 15 N; N mineralization; nitrification; northern Arizona; Pinus ponderosa Laws.; pon- derosa pine-bunchgrass communities; prescribed burning; restoration ecology; tree thinning.

Community-level physiological profiles of bacteria and fungi: Plate type and incubation temperature influences on contrasting soils

Abstract Temperature sensitivity of community-level physiological profiles (CLPPs) was examined for two semiarid soils from the southwestern United States using five different C-substrate profile microtiter plates (Biolog GN2, GP2, ECO, SFN2, and SFP2) incubated at five different temperature regimes. The CLPPs produced from all plate types were relatively unaffected by these contrasting incubation temperature regimes. Our results demonstrate the ability to detect CLPP differences between similar soils with differing physiological parameters, and these differences are relatively insensitive to incubation temperature. Our study also highlights the importance of using both bacterial and fungal plate types when investigating microbial community differences by CLPP. Nevertheless, it is unclear whether or not the differences in CLPPs generated using these plates reflect actual functional differences in the microbial communities from these soils in situ.

From Genes to Ecosystems : The Genetic Basis of Condensed Tannins and Their Role in Nutrient Regulation in a Populus Model System

Research that connects ecosystem processes to genetic mechanisms has recently gained significant ground, yet actual studies that span the levels of organization from genes to ecosystems are extraordinarily rare. Utilizing foundation species from the genus Populus, in which the role of condensed tannins (CT) has been investigated aboveground, belowground, and in adjacent streams, we examine the diverse mechanisms for the expression of CT and the ecological consequences of CT for forests and streams. The wealth of data from this genus highlights the importance of form and function of CT in large-scale and long-term ecosystem processes and demonstrates the following four patterns: (1) plant-specific concentration of CT varies as much as fourfold among species and individual genotypes; (2) large within-plant variation in CT occurs due to ontogenetic stages (that is, juvenile and mature), tissue types (that is, leaves versus twigs) and phenotypic plasticity in response to the environment; (3) CT have little consistent effect on plant–herbivore interactions, excepting organisms utilizing woody tissues (that is, fungal endophytes and beaver), however; (4) CT in plants consistently slow rates of leaf litter decomposition (aquatic and terrestrial), alter the composition of heterotrophic soil communities (and some aquatic communities) and reduce nutrient availability in terrestrial ecosystems. Taken together, these data suggest that CT may play an underappreciated adaptive role in regulating nutrient dynamics in ecosystems. These results also demonstrate that a holistic perspective from genes-to-ecosystems is a powerful approach for elucidating complex ecological interactions and their evolutionary implications.

Merging aquatic and terrestrial perspectives of nutrient biogeochemistry.

Although biogeochemistry is an integrative discipline, terrestrial and aquatic subdisciplines have developed somewhat independently of each other. Physical and biological differences between aquatic and terrestrial ecosystems explain this history. In both aquatic and terrestrial biogeochemistry, key questions and concepts arise from a focus on nutrient limitation, ecosystem nutrient retention, and controls of nutrient transformations. Current understanding is captured in conceptual models for different ecosystem types, which share some features and diverge in other ways. Distinctiveness of subdisciplines has been appropriate in some respects and has fostered important advances in theory. On the other hand, lack of integration between aquatic and terrestrial biogeochemistry limits our ability to deal with biogeochemical phenomena across large landscapes in which connections between terrestrial and aquatic elements are important. Separation of the two approaches also has not served attempts to scale up or to estimate fluxes from large areas based on plot measurements. Understanding connectivity between the two system types and scaling up biogeochemical information will rely on coupled hydrologic and ecological models, and may be critical for addressing environmental problems associated with locally, regionally, and globally altered biogeochemical cycles.

Influence of red alder on soil nitrogen transformations in two conifer forests of contrasting productivity

Abstract We conducted laboratory studies to determine the effects of red alder ( Alnus rubra Bong.) on soil N transformations and N availability indices at two conifer forest sites of contrasting productivity. The inclusion of red alder in conifer forests significantly increased gross rates of N mineralization, N immobilization, nitrification and NO 3 − immobilization, and the effects of alder were generally similar for soils from low- and high-productivity sites. However, the addition of alder to the conifer stand at the high productivity site increased gross N mineralization and immobilization processes more than at the low productivity site. At both sites, gross N and NO 3 − production were enhanced by alder more than gross N immobilization processes, leading to higher rates of net N mineralization and nitrification. At the fertile site, most microbial N assimilation occurred from the NO 3 − pool, compared with less than half at the infertile site (none as NO 3 − in the less productive pure conifer stand). Heterotrophic nitrification (as indicated by a lack of C 2 H 2 inhibition) accounted for 65–72% of the gross nitrification in all stands that exhibited nitrification (no nitrification was detected in the pure conifer stand at the infertile site). The inclusion of red alder had no effect on the proportion of total nitrification that was heterotrophic, despite the lower soil pH in mixed alder-conifer stands compared to conifer stands. Gross rates of N mineralization correlated well with both autotrophic and heterotrophic nitrification across all soils. Gross N mineralization may be a good index of NH 4 + availability to autotrophic nitrifiers, as well as the quality of organic N as a substrate for heterotrophic nitrification. Most estimates of microbial biomass and activity, N availability and N transformation rates were significantly correlated with each other. In general, gross N transformations were better correlated with other indices of N availability and microbial activity than estimates of net N transformations. Similar N cycling rates and microbial biomass N pool sizes in pure alder and adjacent alder-conifer stands at the fertile site suggest that continued inputs of N via symbiotic N-fixation by red alder in coniferous forest stands can lead to the elimination of N-limitation to forest ecosystem production.