Seasonal variation in biomass and carbohydrate partitioning of understory sugar maple (Acer saccharum) and yellow birch (Betula alleghaniensis) seedlings

We investigated seasonal patterns of biomass and carbohydrate partitioning in relation to shoot growth phenology in two age classes of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britt.) seedlings growing in the understory of a partially harvested forest. The high root:shoot biomass ratio and carbohydrate concentration of sugar maple are characteristic of species with truncated growth patterns (i.e., cessation of aboveground shoot growth early in the growing season), a conservative growth strategy and high shade tolerance. The low root:shoot biomass ratio and carbohydrate concentration of yellow birch are characteristic of species with continuous growth patterns, an opportunistic growth strategy and low shade tolerance. In both species, starch represented up to 95% of total nonstructural carbohydrates and was mainly found in the roots. Contrary to our hypothesis, interspecific differences in shoot growth phenology (i.e., continuous versus truncated) did not result in differences in seasonal patterns of carbohydrate partitioning. Our results help explain the niche differentiation between sugar maple and yellow birch in temperate, deciduous understory forests.     

Interspecific variation in nighttime transpiration and stomatal conductance in a mixed New England deciduous forest

Transpiration is generally assumed to be insignificant at night when stomata close in response to the lack of photosynthetically active radiation. However, there is increasing evidence that the stomata of some species remain open at night, which would allow for nighttime transpiration if there were a sufficient environmental driving force. We examined nighttime water use in co-occurring species in a mixed deciduous stand at Harvard Forest, MA, using whole-tree and leaf-level measurements. Diurnal whole-tree water use was monitored continuously with Granier-style sap flux sensors in paper birch (Betula papyrifera Marsh.), red oak (Quercus rubra L.) and red maple (Acer rubrum L.). An analysis was conducted in which nighttime water flux could be partitioned between refilling of internal water stores and transpiration. Substantial nighttime sap flux was observed in all species and much of this flux was attributed to the refilling of depleted water stores. However, in paper birch, nighttime sap flux frequently exceeded recharge estimates. Over 10% of the total daily sap flux during the growing season was due to transpiration at night in paper birch. Nighttime sap flux was over 8% of the total daily flux in red oak and 2% in red maple; however, this flux was mainly associated with recharge. On nights with elevated vapor pressure deficit, sap flux continued through the night in paper birch, whereas it reached zero during the night in red oak and red maple. Measurements of leaf-level gas exchange on a night with elevated vapor pressure deficit showed stomatal conductance dropping by only 25% in paper birch, while approaching zero in red oak and red maple. The study highlighted differences in ecophysiological controls on sap flux exerted by co-occurring species. Paper birch is a fast-growing, shade-intolerant species with an earlier successional status than red oak and red maple. Risking water loss through nighttime transpiration may provide paper birch with an ecological advantage by enabling the species to maximize photosynthesis and support rapid growth. Nighttime transpiration may also be a mechanism for delivering oxygen to respiring cells in the deep sapwood of paper birch.     

Forest regeneration 50 years following partial cutting in mixedwood ecosystems of southern Quebec,Canada

The impact of winter harvesting on regeneration 50 years after an experimental diameter-limit cutting was examined in mixed deciduous–coniferous ecosystems of southern Quebec, Canada. The study was conducted in La Mauricie National Park, Quebec, Canada. Regeneration data in two balsam fir (Abies balsamea (L.) Mill.), red spruce (Picea rubens Sarg.), sugar maple (Acer saccharum Marsh.), and yellow birch (Betula alleghaniensis Britt.) ecosystem types were analyzed. Comparisons between uncut and cut stands were obtained from a total of 63 sample plots. For both ecosystems, there were no significant differences between uncut and cut plots for regeneration density and stocking. The most abundant regeneration species were balsam fir, red spruce, sugar maple, red maple (Acer rubrum L.), yellow birch and American beech (Fagus grandifolia Ehrh.). The type of diameter-limit cutting described in the study did not affect regeneration density and stocking but its impact on productivity, timber quality and genetics is still unknown.     

Development and recovery from winter embolism in silver birch: seasonal patterns and relationships with the phenological cycle in oceanic Scotland

Silver birch (Betula pendula Roth) is increasingly used in the United Kingdom for reforestation. However, recent evidence indicates that, under some circumstances, planted birch can suffer serious and repeated mortality of the apical leaders and branches, with consequent loss of apical dominance and the formation of a contorted stem. Plants from 37 seed sources of silver birch from Scotland and northern England planted at two sites were compared for several characteristics related to hydraulic architecture, vulnerability to freeze-thaw cycle induced embolism and spring recovery from winter embolism during the period 2000-2002. Phenological rhythms were also monitored in late winter-early spring to document relationships between phenology and water relations parameters. Significant differences were found across seed sources in stage of bud flushing for four dates in spring. Early flushing seed sources differed by about 1 to 2 weeks from late-flushing seed sources across the two sites. Wintertime xylem embolism in stems reached a peak of about 50 to 70% loss of xylem hydraulic conductivity, depending on the size and position of the sample shoots in the canopy. Small apical shoots were significantly more embolized than large basal shoots. Development of winter embolism was coupled to the occurrence of frost events. As percent loss of hydraulic conductivity increased during the winter, wood relative water content declined. Embolism reversal occurred rapidly in spring at the time of development of positive root pressure. No significant differences in the degree of winter embolism in 2001 were found among the three seed sources examined. The investigation was expanded in the winter-spring of 2002 to include 10 seed sources across both sites. Significant differences were found in degree of winter embolism across sites, dates and seed sources. For each date, there was a significant relationship between flushing scores and wood relative water contents across the two sites and all seed sources, suggesting that differences in time of flushing across sites and seed sources were likely caused by differences in the time of occurrence of root pressure, a necessary precondition to flushing.     

Study of landscape change under forest harvesting and climate warming-induced fire disturbance

We examined tree species responses under forest harvesting and an increased fire disturbance scenario due to climate warming in northern Wisconsin where northern hardwood and boreal forests are currently predominant. Individual species response at the ecosystem scale was simulated with a gap model, which integrates soil, climate and species data, stratified by ecoregions. Such responses were quantified as species establishment coefficients. These coefficients were used to parameterize a spatially explicit landscape model, LANDIS. Species response to climate warming at the landscape scale was simulated with LANDIS, which integrates ecosystem dynamics with spatial processes including seed dispersal, fire disturbance, and forest harvesting. Under a 5 °C annual temperature increase predicted by global climate models (GCM), our simulation results suggest that significant change in species composition and abundance could occur in the two ecoregions in the study area. In the glacial lake plain (lakeshore) ecoregion under warming conditions, boreal and northern hardwood species such as red oak, sugar maple, white pine, balsam fir, paper birch, yellow birch, and aspen decline gradually during and after climate warming. Southern species such as white ash, hickory, bur oak, black oak, and white oak, which are present in minor amounts before the warming, increase in abundance on the landscape. The transition of the northern hardwood and boreal forest to one dominated by southern species occurs around year 200. In the sand barrens ecoregion under warming conditions, red pine initially benefits from the decline of other northern hardwood species, and its abundance quickly increases. However, red pine and jack pine as well as new southern species are unable to reproduce, and the ecoregion could transform into a region with only grass and shrub species around 250 years under warming climate. Increased fire frequency can accelerate the decline of shade-tolerant species such as balsam fir and sugar maple and accelerate the northward migration of southern species. Forest harvesting accelerated the decline of northern hardwood and boreal tree species. This is especially obvious on the barrens ecoregion, where the intensive cutting regime contributed to the decline of red pine and jack pine already under stressed environments. Forest managers may instead consider a conservative cutting plan or protective management scenarios with limited forest harvesting. This could prolong the transformation of the barrens into prairie from one-half to one tree life cycle.     

Forest regeneration composition and development in upland, mixed-oak forests

Advance regeneration in 52 mature mixed-oak stands was analyzed and described. Red maple (Acer rubrum L.) was the most abundant species in the study area. Among oak (Quercus) species, northern red oak (Q. rubra L.) was the most abundant within the Allegheny Plateau physiographic province, whereas chestnut oak (Q. montana L.) was the most abundant within the Ridge and Valley physiographic province. Sixteen stands, for which data are available through the fourth growing season following harvest, were used to describe stand development. Cumulative height, a composite measure of size and density, was used to describe early stand development. Black gum (Nyssa sylvatica Marsh.) and black birch (Betula lenta L.) had dramatic increases in stand density and cumulative height after overstory removal. Cumulative height of northern red oak and chestnut oak showed a faster positive response to overstory removal than red maple. Oak retained its dominance in cumulative height for at least 4 years after harvest. Red maple nevertheless remained the most abundant tree species after overstory removal. Our results suggest that the principal advantage of red maple regeneration is its ability to accumulate in large numbers prior to harvest.     

Winter embolism,mechanisms of xylem hydraulic conductivity recovery and springtime growth patterns in walnut and peach trees

Xylem vessels of Prunus persica Batsch (peach) and Juglans regia L. (walnut) are vulnerable to frost-induced embolism. In peach, xylem embolism increased progressively over the winter, reaching a maximum of 85% loss of hydraulic conductivity (PLC) in early March. Over winter, PLC in walnut approached 100%, but the degree of xylem embolism varied during the winter, reflecting the ability of walnut to generate positive xylem pressures in winter and spring. In contrast, positive xylem pressures were not observed in peach. Controlled freeze-thaw experiments showed that frost alone is insufficient to increase embolism in peach; evaporative conditions during thawing are also required. However, when both species were protected from frost, PLC was zero. At bud break, there was complete recovery from embolism in walnut, whereas PLC remained high in peach. Three mechanisms responsible for the restoration of branch hydraulic conductivity were identified in walnut: the development of stem pressure, the development of root pressure and the formation of a new ring of functional xylem, whereas only one mechanism was observed in peach (new functional ring). The climatic conditions necessary for the manifestation of these mechanisms were investigated.     

Effects of simulated thaw on xylem cavitation,residual embolism,spring dieback and shoot growth in yellow birch

Yellow birch seedlings (Betula alleghaniensis Britt.) that had lost more than 90% of their stem hydraulic conductivity during ambient winter temperatures were exposed to 0 and 20 days of a simulated winter thaw followed by a 48-h freezing treatment at 0, -5, -10, -20 and -30 degrees C. After measuring freezing injury to shoots and roots, the seedlings were placed in a greenhouse where recovery of xylem conductivity and new growth were measured. Shoot xylem cavitation was measured as percent loss of hydraulic conductivity. Shoot freezing injury was assessed by electrolyte leakage (EL) and root freezing injury was assessed by EL and triphenyl tetrazolium chloride reduction. Seedlings pretreated with thaw had higher stem water contents and suffered more freezing damage to roots and shoots (at -20 and -30 degrees C, respectively) than unthawed seedlings. After 3 weeks in a greenhouse, seedlings from the 0, -5 and -10 degrees C freezing treatments showed complete recovery of xylem conductivity, with substantially increased stem water contents. Poor recovery of hydraulic conductivity was observed only in seedlings that were subjected to freezing treatments at -20 and -30 degrees C, regardless of thaw treatment. Of these embolized seedlings, however, only those not previously thawed showed recovery of hydraulic conductivity or regained stem water content after 9 weeks in the greenhouse. Shoot dieback, bud burst and length of new shoots were significantly related to the extent of stem xylem cavitation and freezing injury. We conclude that (1) the simulated winter thaw predisposed yellow birch seedlings to freezing damage in shoots and roots by dehardening tissues and increasing their water content; (2) root freezing damage in turn affected the seedlings' ability to refill embolized stem xylem, resulting in considerable residual xylem embolism after spring refilling; (3) further recovery of stem xylem conductivity was attributable to growth of new vessels; (4) and the permanent residual embolism, together with root and shoot freezing injury, caused increased dieback, bud mortality and reduced growth of new shoots.     

Variations in amounts of carbohydrates, amino acids and adenine nucleotides in mulberry tree (Morus alba L.) stems during transitional phases of growth

Quantitative changes in carbohydrates, amino acids and adenine nucleotides in the stems of mulberry trees (Morus alba L., cv. Shin-ichinose) were followed from spring to early summer and from autumn to early spring. Both ATP and ADP content of stems increased before bud break, whereas the content of sucrose, the most abundant sugar among the stem carbohydrates, decreased. The sucrose content fell to its lowest value at the beginning of May, and then increased rapidly, whereas the starch content decreased suggesting consumption of reserve carbohydrate and simultaneous accumulation of current photosynthate. This was confirmed by studies in which reserve carbohydrates were labeled with (14)CO(2). Proline content of stems increased from the time of leaf shedding until early spring. Although it was the most abundant amino acid at the time of bud break, proline rapidly decreased as new shoots developed and was hardly detectable by the beginning of May. The asparagine and arginine contents increased transiently following bud break, and then decreased toward summer. Transient increases in glutamine and arginine were noted at the time of leaf shedding.     

Species composition and dynamics in two hardwood stands in Vermont: a disturbance history

Stand composition and structure utilizing stem analysis was studied in two hardwood stands in Vermont. In a mixed hardwood stand with some white pine and hemlock, a major entry of new trees in the main canopy seems associated with harvesting coincident with land exchange. More recent partial cuttings have promoted establishment of new seedlings or development of suppressed advanced-growth shade-tolerant beech, (Fagus grandifolia, Ehrh.), hophornbeam (Ostrya virginiana, (Mill) K. Koch), and striped maple (Acer pensylvanicum, L.). Very few sugar maple and red oak seedlings and saplings are present.

In a northern hardwood stand some red spruce (Picea rubens, Sarg.), that were 240 to 306 years old, became established before any known harvest, and exhibited release following harvests of the mid-1800s. This major harvest, coupled with the differential growth between spruce and hardwoods, and seed/seedling availability, resulted in a major change in stand composition. Trees now in the main canopy of sampled stands appear to have either been released or newly established following various harvests. Harvests have been of such frequency that natural disturbances seem insignificant. Many of the competitive understory species have become abundant following harvests of the 1960s and 1980s and may have been present as advanced growth and responded to the release. Following the harvest of 1981–1982, abundant yellow birch became established on skid trails. Elsewhere in the stand, yellow birch seedlings and saplings are only in great abundance in areas that were possibly sizable gaps following earlier harvesting.

The dynamics of tree entry and growth in gaps of small or large size probably occur in a similar way in many other stands of the region. Though the sampling of this study is limited, there is no suggestion of continuous tree establishment at any particular location, the new age classes seem associated with either a gap or stand replacing disturbance attributed to harvesting.     

Influence of overstory density on ecophysiology of red oak (Quercus rubra) and sugar maple (Acer saccharum) seedlings in central Ontario shelterwoods

A field experiment was established in a second-growth hardwood forest dominated by red oak (Quercus rubra L.) to examine the effects of shelterwood overstory density on leaf gas exchange and seedling water status of planted red oak, naturally regenerated red oak and sugar maple (Acer saccharum Marsh.) seedlings during the first growing season following harvest. Canopy cover of uncut control stands and moderate and light shelterwoods averaged 97, 80 and 49%, respectively. Understory light and vapor pressure deficit (VPD) strongly influenced gas exchange responses to overstory reduction. Increased irradiance beneath the shelterwoods significantly increased net photosynthesis (P(n)) and leaf conductance to water vapor (G(wv)) of red oak and maple seedlings; however, P(n) and G(wv) of planted and naturally regenerated red oak seedlings were two to three times higher than those of sugar maple seedlings in both partial harvest treatments, due in large part to decreased stomatal limitation of gas exchange in red oak as a result of increased VPD in the shelterwoods. In both species, seedling water status was higher in the partial harvest treatments, as reflected by the higher predawn leaf water potential and seedling water-use efficiency in seedlings in shelterwoods than in uncut stands. Within a treatment, planted and natural red oak seedlings exhibited similar leaf gas exchange rates and water status, indicating little adverse physiological effect of transplanting. We conclude that the use of shelterwoods favors photosynthetic potential of red oak over sugar maple, and should improve red oak regeneration in Ontario.     

Investigation of change in tangential strain on the inner bark of the stem and root ofBetula platyphylla var.japonica andAcer mono during sap season

Although it is well known that sap exudation during early spring in temperate deciduous trees occurs in response to daytime warming and nighttime cooling, the mechanisms of the process are not yet fully understood. Previous theories suggested that changes in stress in the wood caused by daytime heating and nighttime cooling might be linked with sap flow. Consequently, a study of itaya-kaede maple (Aver mono) and shirakamba birch (Betula platyphylla var.japonica) looked at tangential strains. One-hour intervals for 3 years of the tangential strains on the inner bark of stem and root were measured in itaya-kaede maple and shirakamba birch during the sap exudation season. The measurements indicated different mechanisms of sap exudation in these two trees. During the sap exudation season in late March, when the temperature fluctuated around 0°C, the tangential strain in the root of itaya-kaede maple showed expansion in the daytime and contraction at night. Conversely, in early April the tangential strain in the root of shirakamba birch exhibited contraction in the daytime and expansion at night. The changes in tangential strains in itaya-kaede maple were attributed to conditioning, a known concept used to explain the uptake mechanism of soil water in maple and its exudation during early spring. However, because the change in tangential strain in the roots of shirakamba birch was similar to that found during the rampant season, sap exudation was not attributed to conditioning but to the plentiful supply of water from the roots. The implications of these mechanisms are that different sap harvesting techniques may be appropriate for different tree species.     

Fall frost resistance in willows used for biomass production. II. Predictive relationships with sugar concentration and dry matter content

The accumulation of sugars and dry matter in stems in fall was examined in relation to frost hardening in eight willow clones (six clones of Salix viminalis L. and one clone each of S. viminalis x S. schwerenii E. Wolf and S. dasyclados Wimm.). Evidence is presented that three sources of variation in fall frost resistance among the eight clones could be assessed from an analysis of stem composition. First, the pre-hardening value of frost resistance could be assessed from the total sugar concentration. Second, the start of induction of apical growth cessation and hence frost hardening could be distinguished by a stepwise increase in sucrose-to-glucose ratio. Third, the progress of frost hardening during its first phase could be followed from a proportional rise in total sugar concentration and, even more accurately, from a proportional rise in dry-to-fresh weight ratio. In contrast, the second phase of frost hardening was largely uncoupled from sugar and dry matter accumulation. Raffinose and sucrose accumulation seemed to be under differential environmental controls. Sucrose accumulation started with the initiation of growth cessation controlled by photoperiod, whereas raffinose accumulation started with falling temperatures later on. Starch reserves that built up in stems in early fall were partially mobilized later on to support sugar accumulation. In contrast to stems, leaves did not exhibit a preferential accumulation of sucrose in fall.     

Acclimation of shade-developed leaves on saplings exposed to late-season canopy gaps

We hypothesized that photoinhibition of shade-developed leaves of deciduous hardwood saplings would limit their ability to acclimate photosynthetically to increased irradiance, and we predicted that shade-tolerant sugar maple (Acer saccharum Marsh.) would be more susceptible to photoinhibition than intermediately shade-tolerant red oak (Quercus rubra L.). After four weeks in a canopy gap, photosynthetic rates of shade-developed leaves of both species had increased in response to the increase in irradiance, although final acclimation was more complete in red oak. However, photoinhibition occurred in both species, as indicated by short-term reductions in maximum rates of net photosynthesis and the quantum yield of oxygen evolution, and longer-term reductions in the efficiency of excitation energy capture by open photosystem II (PSII) reaction centers (dark-adapted F(v)/F(m)) and the quantum yield of PSII in the light (phi(PSII)). The magnitude and duration of this decrease were greater in sugar maple than in red oak, suggesting greater susceptibility to photoinhibition in sugar maple. Photoinhibition may have resulted from photodamage, but it may also have involved sustained rates of photoprotective energy dissipation (especially in red oak). Photosynthetic acclimation also appeared to be linked to an ability to increase leaf nitrogen content. Limited photosynthetic acclimation in shade-developed sugar maple leaves may reflect a trade-off between shade-tolerance and rapid acclimation to a canopy gap.     

Long- and short-term flooding effects on survival and sink-source relationships of swamp-adapted tree species

About 95% of swamp tupelo (Nyssa sylvatica var. biflora (Walt.) Sarg.) and sweetgum (Liquidambar styraciflua L.) seedlings survived continuous root flooding for more than two years, whereas none of the swamp chestnut oak (Quercus michauxii Nutt.) and cherrybark oak (Q. falcata var. pagodifolia Ell.) seedlings survived one year of flooding. Death of oak seedlings occurred in phases associated with periods of major vegetative growth, e.g., after bud burst in spring, after summer stem elongation, and during the winter deciduous stage, suggesting that stored reserves and sources were inadequate to maintain the seedlings when vegetative sinks were forming. Additional evidence that flooding induced a source deficiency in oak was that leaves of flooded oak were 65 to 75% smaller than leaves of nonflooded oak. Flooded swamp tupelo seedlings had a normal leaf size and patchy stomatal opening compared with nonflooded seedlings. Flooding caused increases in alcohol dehydrogenase (ADH) specific activity in taproot cambial tissues and increases in starch concentrations of swamp tupelo seedlings that were reversed when seedlings were removed from flooding. Flooding had little effect on soluble sugar concentrations in swamp tupelo or sweetgum. In the long-term flood-dry-flood treatment, in which all species had survivors, upper canopy leaf photosynthetic rates were higher in all species during the dry period than in nonflooded controls, whereas their starch and soluble sugars concentrations were similar to those of nonflooded controls. Based on seedling survival and the sink-source relationships, the order of flood tolerance was: swamp tupelo > sweetgum > swamp chestnut oak > cherrybark oak.     

Fine root dynamics of oak saplings in response to Phytophthora cinnamomi infection under different temperatures and durations

The belowground effects of Phytophthora cinnamomi on 1‐year‐old saplings of two common oak species in mid‐Atlantic US forests, white (Quercus alba) and red oak (Q. rubra), were examined after incubation in pathogen‐infested soilless potting mix. Fine root lengths (0–1.5 mm in diameter) of both oak species were quantified after incubation at successive 30‐day intervals up to 300 days, for a total of 10 incubation periods. In addition, colony‐forming units (CFU) of P. cinnamomi were quantified after white oak saplings were incubated in infested soilless potting mix at different temperature/duration combinations that reflect soil conditions present in the mid‐Atlantic United States. Impact of P. cinnamomi on fine root lengths of red and white oak saplings varied considerably over time. Significant periods of fine root loss occurred primarily during spring, when bud break and leaf flush began for both oak species. Red oaks had 17% fine root loss on average, while white oaks appeared more resistant to P. cinnamomi infection with a 2% decrease in fine roots over the course of the experiment. Phytophthora cinnamomi CFU declined significantly with exposure to all incubation temperatures except 8°C. This was in contrast to in vitro experiments, where the optimum temperature for mycelial growth was determined to be 21°C and above. Significant fine root loss caused by P. cinnamomi depended on plant phenology and the oak species tested. Extreme soil temperatures have a significant adverse impact on temporal changes of P. cinnamomi population.     

Dendroecology and species co-existence in an old-growth Quercus–Acer–Tilia talus slope forest in the central Appalachians,USA

Dendroecological techniques were used to examine the disturbance history and patterns of species recruitment in an old-growth Quercus rubra L. (northern red oak)–Acer saccharum Marsh. (sugar maple)–Tilia americana L. (basswood) forest on a steep, talus slope in eastern West Virginia. The forest was uneven-aged as were the populations of red oak. Sugar maple dominated the sapling layer, which comprised little or no basswood and red oak. A compilation of major and moderate releases (indicative of disturbance) in 25 cores revealed single or multiple release events in every decade from 1870–1990. The high elevation of the forest coupled with a fertile sub-soil beneath the talus ameliorated the outwardly harsh conditions of the site, allowing for the domination of typically mesophytic, nutrient demanding tree species. We observed several fire scarred trees as well as extensive small-scale blow-down throughout the forest. Frequent disturbance events were probably crucial to the co-existence and continuous canopy recruitment of the relatively light demanding red oak with highly shade tolerant sugar maple and basswood. The strong successional replacement tendencies of red oak by northern hardwoods noted elsewhere in the eastern US may be less apparent on high elevation, rocky sites in the central Appalachians. Thus, this is a unique case study of long-term red oak domination with later successional species in an old-growth forest.     

Water use and carbon exchange of red oak- and eastern hemlock-dominated forests in the northeastern USA: implications for ecosystem-level effects of hemlock woolly adelgid

Water use and carbon exchange of a red oak-dominated (Quercus rubra L.) forest and an eastern hemlock-dominated (Tsuga canadensis L.) forest, each located within the Harvard Forest in north-central Massachusetts, were measured for 2 years by the eddy flux method. Water use by the red oak forest reached 4 mm day(-1), compared to a maximum of 2 mm day(-1) by the eastern hemlock forest. Maximal carbon (C) uptake rate was also higher in the red oak forest than in the eastern hemlock forest (about 25 versus 15 micromol m(-2) s(-1)). Sap flux measurements indicated that transpiration of red oak, and also of black birch (Betula lenta L.), which frequently replaces eastern hemlock killed by hemlock woolly adelgid (Adelges tsugae Annand.), were almost twice that of eastern hemlock. Despite the difference between species in maximum summertime C assimilation rate, annual C storage of the eastern hemlock forest almost equaled that of the red oak forest because of net C uptake by eastern hemlock during unusually warm fall and spring weather, and a near-zero C balance during the winter. Thus, the effect on C storage of replacing eastern hemlock forest with a forest dominated by deciduous species is unclear. Carbon storage by eastern hemlock forests during fall, winter and spring is likely to increase in the event of climate warming, although this may be offset by C loss during hotter summers. Our results indicate that, although forest water use will decrease immediately following eastern hemlock mortality due to the hemlock woolly adelgid, the replacement of eastern hemlock by deciduous species such as red oak will likely increase summertime water use over current rates in areas where hemlock is a major forest species.     

Element accumulation patterns of deciduous and evergreen tree seedlings on acid soils: implications for sensitivity to manganese toxicity

Foliar nutrient imbalances, including the hyperaccumulation of manganese (Mn), are correlated with symptoms of declining health in sensitive tree species growing on acidic forest soils. The objectives of this study were to: (1) compare foliar nutrient accumulation patterns of six deciduous (sugar maple (Acer saccharum Marsh.), red maple (Acer rubrum L.), red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.) and white ash (Fraxinus americana L.)) and three evergreen (eastern hemlock (Tsuga canadensis L.), white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss.)) tree species growing on acidic forest soils; and (2) examine how leaf phenology and other traits that distinguish evergreen and deciduous tree species influence foliar Mn accumulation rates and sensitivity to excess Mn. For the first objective, leaf samples of seedlings from five acidic, non-glaciated field sites on Pennsylvania's Allegheny Plateau were collected and analyzed for leaf element concentrations. In a second study, we examined growth and photosynthetic responses of seedlings exposed to excess Mn in sand culture. In field samples, Mn in deciduous foliage hyperaccumulated to concentrations more than twice as high as those found in evergreen needles. Among species, sugar maple was the most sensitive to excess Mn based on growth and photosynthetic measurements. Photosynthesis in red maple and red oak was also sensitive to excess Mn, whereas white oak, black cherry, white ash and the three evergreen species were tolerant of excess Mn. Among the nine species, relative rates of photosynthesis were negatively correlated with foliar Mn concentrations, suggesting that photosynthetic sensitivity to Mn is a function of its rate of accumulation in seedling foliage.