ALBERT

Description: We surveyed and wounded forest-grown sugar maple ( Acer saccharum Marsh.) trees in a long-term, replicated Ca manipulation study at the Hubbard Brook Experimental Forest in New Hampshire, USA. Plots received applications of Ca (to boost Ca availability above depleted ambient levels) or Al (to compete with Ca uptake and further reduce Ca availability). We found significantly greater total foliar and membrane-associated Ca in foliage of trees in plots fertilized with Ca when compared with trees from Al-addition and control plots (P = 0.005). Coinciding with foliar Ca differences, trees exhibited a significant difference in crown vigor and in percent branch dieback among treatments (P 〈 0.05), with a trend towards improved canopy health as Ca levels increased. Annual basal area increment growth for the years following treatment initiation (1998–2004) was significantly greater in trees subjected to Ca addition compared with trees in control and Al treatments. Treatment-related improvements in growth were particularly evident after overstory release following a 1998 ice storm. The amount of wound closure was also greatest for trees in Ca-addition plots relative to Al-addition and control plots (P = 0.041). These findings support evidence that ambient Ca depletion is an important limiting factor regarding sugar maple health and highlight the influence of Ca on wound closure and growth following release from competition.

Description: Techniques are described for field and laboratory extraction of small quantities of sugar maple (Acersaccharum Marsh.) xylem sap when pressure in the xylem is less than atmospheric. Accurate estimates of sap sugar concentration can be made with a hand refractometer most of the year. There is, however, appreciable within-tree variation in sap sugar concentration at any given time and within short periods of time.

Description: Autumnal leaf anthocyanin expression is enhanced following exposure to a variety of environmental stresses and may represent an adaptive benefit of protecting leaves from those stresses, thereby allowing for prolonged sugar and nutrient resorption. Past work has shown that experimentally induced sugar accumulations following branch girdling triggers anthocyanin biosynthesis. We hypothesized that reduced phloem transport at low autumnal temperatures may increase leaf sugar concentrations that stimulate anthocyanin production, resulting in enhanced tree- and landscape-scale color change. We used refrigerant-filled tubing to cool individual branches in a mature sugar maple (Acer saccharum Marsh.) tree to test whether phloem cooling would trigger foliar sugar accumulations and enhance anthocyanin biosynthesis. Cooling increased foliar sucrose, glucose, and fructose concentrations 2- to nearly 10-fold (depending on the specific sugar and sampling date) relative to controls and increased anthocyanin concentrations by approximately the same amount. Correlation analyses indicated a strong and steady positive relationship between anthocyanin and sugar concentrations, which was consistent with a mechanistic link between cooling-induced changes in these constituents. Tested here at the branch level, we propose that low temperature induced reductions in phloem transport may be responsible for increases in foliar sugars that trigger anthocyanin displays at grander scales.

Description: Red oak (Quercus rubra L.) is projected to expand into the northern hardwood forest over the coming century. We explored the connection between red oak basal area growth and a number of factors (tree age and size, stand dynamics, site elevation, and climate and acid deposition variables) for 213 trees in 11 plots throughout Vermont, USA. Red oak growth generally increased over the course of the chronology (1935–2014) and has been particularly high in recent decades. Growth differed among elevational groups but did not differ between age or size groups. Summer moisture metrics were consistently and positively associated with growth, whereas fall moisture was associated with reduced growth in recent decades. Higher summer temperatures were often negatively associated with growth, though there was evidence that low temperatures in the summer (higher elevations) and fall (lower elevations) constrain growth. Several pollution metrics were associated with reduced growth, a surprising result for a species not known to be sensitive to inputs of acid deposition that have predisposed other species in the region to decline. While red oak growth is currently robust, increases in summer temperatures, reductions in growing season precipitation, or increases in fall precipitation could reduce future growth potential.

Description: Laboratory experiments have verified that acid-deposition-induced calcium (Ca) leaching reduces the foliar cold tolerance of red spruce (Picea rubens Sarg.) current-year foliage, increasing the risk of winter injury and crown deterioration. However, to date no studies have shown that ambient losses in soil Ca have resulted in increased winter injury in the field. In 2003, a year of severe region-wide winter injury to red spruce, we measured the nutrition and winter injury of current-year foliage and bud mortality for red spruce on two watersheds at the Hubbard Brook Experimental Forest in Thornton, New Hampshire: (1) a reference watershed that has undergone considerable Ca loss attributed to acid-deposition-induced leaching and (2) a watershed that was fertilized with CaSiO3 in 1999 to replace lost Ca. For all crown classes combined, winter injury was significantly greater (P = 0.05) for red spruce on the reference watershed than for spruce on the Ca-addition watershed. Differences in foliar injury were particularly evident for dominant and codominant trees. For these crown classes, red spruce on the reference watershed lost about 75% of their current-year foliage to winter injury, about three times more than foliar losses for the Ca-addition watershed (P = 0.01). Patterns of bud mortality followed that of foliar injury. The only difference in foliar cation nutrition detected was a significantly greater concentration of Ca in red spruce foliage from the Ca-addition watershed relative to spruce from the reference watershed (P = 0.001). Differences in Ca concentration, foliar winter injury, and bud mortality that occurred coincident with watershed Ca treatment provide the first evidence that ambient Ca depletion is associated with elevated winter injury of red spruce trees.

Description: Abundant winter injury to the current-year (2002) foliage of red spruce (Picea rubens Sarg.) became apparent in the northeastern United States in late winter of 2003. To assess the severity and extent of this damage, we measured foliar winter injury at 28 locations in Vermont and surrounding states and bud mortality at a subset of these sites. Ninety percent of all trees assessed showed some winter injury, and trees lost an average of 46% of all current-year foliage. An average of 32% of buds formed in 2002 were killed in association with winter injury. Both foliar and bud mortality increased with elevation and with crown dominance, and bud mortality increased with greater foliar injury. Foliar injury in 2003 at a plantation near Colebrook, New Hampshire, was more than five times the typical levels for 9 previous years of measurement and more than twice that measured for another high-injury year. Plantation data also indicated that bud mortality in 2003 was greater than previously documented and that persistent winter injury was associated with increased tree mortality. Comparisons of our data with past studies for two sites with native red spruce also indicated that damage in 2003 was greater than other recently reported, high-injury years. Because heavy foliar and bud losses can severely disrupt the carbon economies of trees, the 2003 winter injury event could lead to further spruce decline and mortality, particularly among dominant trees at higher elevations.

Description: Red spruce (Picea rubens Sarg.) winter injury is caused by freezing damage that results in the abscission of the most recent foliar age-class. Injury was widespread and severe in the northeastern United States in 2003 and was assessed at multiple elevations at 23 sites in Vermont and adjacent states. This paper presents a spatial analysis of these injury assessments. Relationships between winter injury on dominant and codominant spruce trees and elevation, latitude, longitude, slope, and aspect were investigated with least squares regression and geographically weighted regression. Results of these analyses indicate that injury increased (1) with elevation; (2) from east to west; (3) with the degree to which plots faced west, except at the highest elevations, where injury was uniformly severe; (4) with increases in slope steepness at higher elevations, or with decreases in slope steepness at lower elevations; and (5) with the degree to which plots faced south, except at the highest elevations in northern locations, where injury was uniformly severe. Because injury was greater in areas that have historically received higher levels of acid and nitrogen deposition western portions of the study region, west-facing slopes, and higher elevations observed patterns of injury support the hypothesis that acidic and (or) nitrogen deposition act on a landscape scale to exacerbate winter injury. Greater injury on south-facing slopes suggests that sun exposure exacerbates injury or its expression.

Description: Acid deposition induced losses of calcium (Ca) from northeastern forests have had negative effects on forest health for decades, including the mobilization of potentially phytotoxic aluminum (Al) from soils. To evaluate the impact of changes in Ca and Al availability on sugar maple (Acer saccharum Marsh.) and American beech (Fagus grandifolia Ehrh.) growth and forest composition following a major ice storm in 1998, we measured xylem annual increment, foliar cation concentrations, American beech root sprouting, and tree mortality at the Hubbard Brook Experimental Forest (Thornton, New Hampshire) in control plots and in plots amended with Ca or Al (treated plots) beginning in 1995. Dominant sugar maple trees were unaffected by the treatment, but nondominant sugar maple tree growth responded positively to Ca treatment. Although plots were mainly composed of sugar maple, American beech experienced the greatest growth on Al-treated plots. Increases in tree mortality on Al-treated plots may have released surviving American beech and increased their growth. The Al tolerance of American beech and the Ca:Al sensitivity of sugar maple contributed to divergent growth patterns that influenced stand productivity and composition. Given that acidic inputs are expected to continue, the growth dynamics associated with Al treatment may have direct relevance to future conditions in native forests.

Description: Despite considerable study, it remains uncertain what environmental factors contribute to red spruce ( Picea rubens Sarg.) foliar winter injury and how much this injury influences tree C stores. We used a long-term record of winter injury in a plantation in New Hampshire and conducted stepwise linear regression analyses with local weather and regional pollution data to determine which parameters helped account for observed injury. Two types of weather phenomena were consistently associated with elevated injury: (i) measures of low-temperature stress that incite injury and (ii) factors that reduced the length of the growing season and predisposed trees to injury. At this plantation, there was a significant linear relationship between winter injury and growth reductions for 2 years after a severe winter injury event. Analysis using data from three New England states indicated that plantation data reflected a regional response. Using regional data, we estimated a reduction of 394 000 metric tons of C sequestered in living red spruce stems ≥20 cm in diameter growing in New York and northern New England during the 2 years following a severe winter injury event. This is a conservative estimate of reduced C sequestration because injury-induced mortality and other factors were not evaluated.

Description: We examined patterns of genetic structuring within a mature eastern white pine (Pinus strobus L.) forest, using geographic information system (GIS)-based data and maps that combined genetic (isozyme analysis of 46 loci) and other tree-specific information (e.g., size, growth, age, and location) for 220 trees in Jericho, Vermont. Interconnections between genotypic information with other tree characteristics revealed several patterns of genetic structuring. Average observed heterozygosity generally increased with tree age-class, and trees with a high number of rare alleles were disproportionally represented in suppressed crown classes. Spatial structuring was also evident: trees within 5 m of one another were highly related, and levels of relatedness generally decreased with increasing distance between trees. In general, a 35-m-radius circle around any tree circumscribed its zone of genetic similarity. Hierarchical cluster analysis indicated the stand consisted of five family groups that exhibited greater genetic similarity within than among clusters. Temporal structuring (a generation gap) was also evident: trees of similar age showed significant positive relatedness, as did trees 3040 years apart. Patterns of genetic structuring likely resulted from the combined influences of natural selection, isolation by distance, and functional generation times. Genetic structuring may also have biological and management implications. Computer-based simulated harvests suggested that the stand could experience genetic alteration when tree removal criteria disrupted existing structural patterns.