Toward the improvement of total nitrogen deposition budgets in the United States.

Frameworks for limiting ecosystem exposure to excess nutrients and acidity require accurate and complete deposition budgets of reactive nitrogen (Nr). While much progress has been made in developing total Nr deposition budgets for the U.S., current budgets remain limited by key data and knowledge gaps. Analysis of National Atmospheric Deposition Program Total Deposition (NADP/TDep) data illustrates several aspects of current Nr deposition that motivate additional research. Averaged across the continental U.S., dry deposition contributes slightly more (55%) to total deposition than wet deposition and is the dominant process (>90%) over broad areas of the Southwest and other arid regions of the West. Lack of dry deposition measurements imposes a reliance on models, resulting in a much higher degree of uncertainty relative to wet deposition which is routinely measured. As nitrogen oxide (NOx) emissions continue to decline, reduced forms of inorganic nitrogen (NHx = NH3 + NH4+) now contribute >50% of total Nr deposition over large areas of the U.S. Expanded monitoring and additional process-level research are needed to better understand NHx deposition, its contribution to total Nr deposition budgets, and the processes by which reduced N deposits to ecosystems. Urban and suburban areas are hotspots where routine monitoring of oxidized and reduced Nr deposition is needed. Finally, deposition budgets have incomplete information about the speciation of atmospheric nitrogen; monitoring networks do not capture important forms of Nr such as organic nitrogen. Building on these themes, we detail the state of the science of Nr deposition budgets in the U.S. and highlight research priorities to improve deposition budgets in terms of monitoring and flux measurements, leaf- to regional-scale modeling, source apportionment, and characterization of deposition trends and patterns.

Ecophysiology of Acer rubrum seedlings from contrasting hydrologic habitats: growth, gas exchange, tissue water relations, abscisic acid and carbon isotope discrimination.

Eight red maple (Acer rubrum L.) provenances, four each from wet and dry sites, were grown under the same conditions and their physiological responses to soil water availability investigated. Under well-watered conditions, seedlings of wet-site provenances grew faster and had consistently higher net photosynthesis, leaf conductance, maximum carboxylation rate, maximum rate of coupled photosynthetic electron transport, apparent quantum use efficiency, light-saturated photosynthesis and dark respiration than seedlings of dry-site provenances. Under conditions of low soil water availability, only dry-site provenances responded with decreased osmotic potential at full hydration and at the turgor loss point; however, provenances from wet sites showed a smaller reduction in absolute growth rate, a greater reduction in gas exchange and a greater increase in abscisic acid concentrations than dry-site provenances.

Responses of Six Eurasian Ulmus Cultivars to a North American Elm Yellows Phytoplasma.

Elms (genus Ulmus) of six clonal cultivars representing Eurasian species and hybrids were grafted when 2 to 3 years old with bark patches from U. rubra infected with an elm yellows phytoplasma or were left untreated as controls. The cultivars were U. glabra × minor 'Pioneer', U. minor × parvifolia 'Frontier', U. parvifolia 'Pathfinder', U. wilsoniana 'Prospector', and the complex hybrids 'Homestead' and 'Patriot'. Trees were evaluated for infection and symptoms 1 or 2 years after inoculation. Infection was detected via the 4',6-diamidino-2-phenylindol e·2HCl (DAPI) fluorescence test in 26 of 86 grafted trees representing five cultivars. Infection of selected trees was confirmed by polymerase chain reaction (PCR) amplification of a fragment of phytoplasmal rDNA, and the phytoplasma was identified by restriction fragment length polymorphism (RFLP) analysis of the amplified DNA using restriction enzymes AluI, RsaI, and TaqI. Elm yellows phytoplasma was also identified by nested PCR and RFLP analysis in two of seven inoculated, healthy-appearing, DAPI-negative trees and one noninoculated control tree. All RFLP profiles were identical to that of reference strain EY1. Phytoplasma-associated symptoms, observed in five cultivars, included suppressed growth, progressive size reduction of apical shoots and leaves, chlorosis, foliar reddening, witches'-brooms, and dieback. Phyto-plasma was not detected in cv. Homestead. Possible resistance of this cultivar to elm yellows phytoplasma was indicated by localized phloem necrosis in stems below inoculum patches.

Responses of 11 Fraxinus Cultivars to Ash Yellows Phytoplasma Strains of Differing Aggressiveness.

Five cultivars of Fraxinus americana (white ash) and five of F. pennsylvanica (green ash) were graft-inoculated with three strains of ash yellows phytoplasmas at Ames, IA, and with thrsee other strains at Ithaca, NY. A sixth green ash cultivar was tested only in New York. Trees were allowed to grow in field plots for 3 years. Infection was detected via the DAPI (4', 6-diamidino-2-phenylindole 2HCl) fluorescence test. Incidence of witches'-brooms on infected trees was greater on white ash than green ash and varied significantly among phytoplasma strain treatments at both locations. Volume growth of infected ash, averaged across cultivars over 2 years in Iowa and 3 years in New York, was 49 and 59%, respectively, as great as that of noninfected trees. Foliar greenness was reduced significantly by infection at both locations, and this reduction was positively correlated with growth reduction. Cultivars at each location varied significantly in growth of noninfected trees and in growth of diseased trees relative to that of nonin-fected trees (a measure of phytoplasma tolerance), but cultivar means for these variables in Iowa were not significantly correlated with those in New York. Green ash cvs. Bergeson, Dakota Centennial, and Patmore and white ash cv. Autumn Applause were above average in tolerance at both locations. Phytoplasma strains at each location varied significantly in aggressiveness as indicated by host growth suppression.

Comparisons of Tolerance of Ash Yellows Phytoplasmas in Fraxinus Species and Rootstock-Scion Combinations.

Growth responses of different ash (Fraxinus) species and rootstock-scion combinations to ash yellows (AshY) phytoplasmas were compared in greenhouse experiments by expressing each measurement as a proportion of the final average value of the variable in noninoculated, own-rooted control trees. Phytoplasmal infection suppressed shoot growth of white ash (F. americana) and green ash (F. pennsylvanica) beginning when buds opened, but did not suppress velvet ash (F. velutina) until after 60 days of growth. AshY-associated growth losses in height, stem diameter, and root volume, averaged across two experiments, were 80, 93, and 98%, respectively, in white ash; 60, 57, and 79% in green ash; and 23, 0, and 12% in velvet ash. Growth in height, but not in stem diameter or root volume, of diseased white ash on velvet ash rootstock was significantly greater than growth of diseased own-rooted white ash. White ash witches'-brooms grafted onto healthy velvet ash continued to grow but did not produce vigorous, dominant shoots. Growth of diseased velvet ash on white ash roots was severely suppressed in comparison with that of diseased own-rooted velvet ash. Management of AshY through the use of tolerant genotypes may require tolerance in both scions and rootstocks.

An improved method for using electrolyte leakage to assess membrane competence in plant tissues.

A new expression for ion leakage from plant tissue, the tissue ionic conductance (g(Ti)), is compared with electrical conductivity (EC) and a commonly used damage index (I(d)) to test the ability of each expression to correctly describe leakiness in two model systems representing examples of physiological processes with well-known effects on membrane permeability. In experiments in which drought-acclimated leaves were compared with nonacclimated leaves and senescing leaves were compared with nonsenescing leaves, I(d) contradicted our expectation that acclimated tissue would be less leaky than nonacclimated tissue, and g(Ti) and EC confirmed this expectation. In a comparison of senescing and nonsenescing tissue, I(d) again contradicted our expectation that senescing tissue would be more leaky than nonsenescing, and EC and g(Ti) were confirming. Using a diffusion analysis approach, we show that I(d) fails to account for variation in the concentration gradient between the tissue and the bathing solution and variation in the surface area through which efflux occurs. Furthermore, because I(d) is a parameter that relates treatment performance to control performance as a percentage value, it distorts the actual differences among treatments. The resulting artifacts lead to a presentation of membrane integrity which is probably incorrect. EC is a more direct measurement of net ion efflux and appears to be less vulnerable to artifact. However, because g(Ti) is the only expression that explicitly includes chemical driving force and tissue surface area, it is the most reliable of the three expressions.

Response of five temperate deciduous tree species to water stress.

Gas exchange, tissue water relations, and leaf/root dry weight ratios were compared among young, container-grown plants of five temperate-zone, deciduous tree species (Acer negundo L., Betula papyrifera Marsh, Malus baccata Borkh, Robinia pseudoacacia L., and Ulmus parvifolia Jacq.) under well-watered and water-stressed conditions. There was a small decrease (mean reduction of 0.22 MPa across species) in the water potential at which turgor was lost (Psi(tlp)) in response to water stress. The Psi(tlp) for water-stressed plants was -1.18, -1.34, -1.61, -1.70, and -2.12 MPa for B. papyrifera, A. negundo, U. parvifolia, R. pseudoacacia, and M. baccata, respectively. Variation in Psi(tlp) resulted primarily from differences in tissue osmotic potential and not tissue elasticity. Rates of net photosynthesis declined in response to water stress. However, despite differences in Psi(tlp), there were no differences in net photosynthesis among water-stressed plants under the conditions of water stress imposed. In A. negundo and M. baccata, water use efficiency (net photosynthesis/transpiration) increased significantly in response to water stress. Comparisons among water-stressed plants showed that water use efficiency for M. baccata was greater than for B. papyrifera or U. parvifolia. There were no significant differences in water use efficiency among B. papyrifera, U. parvifolia, A. negundo, and R. pseudoacacia. Under water-stressed conditions, leaf/root dry weight ratios (an index of transpiration to absorptive capacity) ranged from 0.77 in R. pseudoacacia to 1.05 in B. papyrifera.