Acclimation of leaves to contrasting irradiance in juvenile trees differing in shade tolerance

Leaves developing in different irradiances undergo structural and functional acclimation, although the extent of trait plasticity is species specific. We tested the hypothesis that irradiance-induced plasticity of photosynthetic and anatomical traits is lower in highly shade-tolerant species than in moderately shade-tolerant species. Seedlings of two evergreen conifers, shade-tolerant Abies alba Mill. and moderately shade-tolerant Picea abies Karst., and two deciduous angiosperm species, highly shade-tolerant Fagus sylvatica L. and moderately shade-tolerant Acer pseudoplatanus L., were grown in deep shade (LL, 5% of full irradiance) or in full solar irradiance (HL) during 2003 and 2004. Steady state responses of quantum yield of PSII (Phi(PSII)), apparent electron transport rate (ETR), nonphotochemical quenching (NPQ) and photochemical quenching (qP) were generally modified by the light environment, with slower declines in Phi(PSII) and qP and greater maximal ETR and NPQ values in HL plants in at least one season; however, no link between quantitative measures of plasticity of these traits and shade tolerance was found. Plasticity of nine anatomical traits (including palisade cell length, which was reduced in LL) showed no relationship with shade tolerance, but was less in conifers than in deciduous trees, suggesting that leaf life span may be a significant correlate of plasticity. When LL-acclimated plants were exposed to HL conditions, the degree and duration of photoinhibition (measured as a decline in maximum quantum yield) was greatest in F. sylvatica, much lower in P. abies and A. alba, and lowest in A. pseudoplatanus. Thus, as with the other traits studied, vulnerability to photoinhibition showed no relationship with shade tolerance.     

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.     

Interactive effects of leaf age and self-shading on leaf structure,photosynthetic capacity and chlorophyll fluorescence in the rain forest tree,Dryobalanops aromatica

In the tropical canopy tree, Dryobalanops aromatica Gaertn. f., upper-canopy leaves (UL) develop under sunlit conditions but are subjected to self-shading within the crown as they age. In contrast, lower-canopy leaves (LL) are exposed to uniform dim light conditions throughout their life span. By comparing leaf morphology and physiology of UL and LL, variations in leaf characteristics were related to leaf age and self-shading. Mass-based chlorophyll (chl) concentration and the chlorophyll/nitrogen (chl/N) ratio were lower and the chl a/b ratio was higher in UL than in LL. In UL, the chl/N ratio gradually increased and the chl a/b ratio gradually decreased with leaf aging, whereas these ratios remained unchanged with leaf age in LL. The effective quantum yield of photosystem II (PSII) (DeltaF/F(m)') at a given irradiance remained unchanged with leaf age in LL, whereas DeltaF/F(m)' changed with leaf age in UL. These data indicate N reallocation within the leaves from carbon fixation components to light harvesting components and a dynamic regulation of photochemical processes of PSII in response to increased self-shading of UL. Despite the difference in light environment with leaf age between UL and LL, maximum photosynthetic rates and nitrogen-use efficiency decreased with leaf aging in both UL and LL. Constancy in the chl/N ratio with leaf age in LL indicated that the decrease in photosynthetic capacity was caused by effects other than shading, such as leaf aging. We conclude that N reallocation and acclimation of PSII to self-shading occurred even in mature leaves, whereas the change in photosynthetic capacity with leaf age was more conservative.     

Leaf-level acclimation to gap creation in mature Acer saccharum trees

Leaf-level morphological and physiological responses of mature, winter-deciduous, shade-tolerant Acer saccharum Marsh. trees to gap formation caused by selection harvest were studied experimentally over a 2-year period. We found no evidence for either physiological stress or positive acclimation following gap creation during the 1-2-week post-harvest period. Rather, lower-canopy leaves showed gradual increases in area-based maximum photosynthetic rates (Amax-area), stomatal conductance (gs), and leaf nitrogen concentration (Narea) over the entire 2-year study. These acclimation responses were directly related to changes in leaf mass per unit area (LMA) in the subsequent two leaf flushes. No change in Amax-area, gs, Narea, or photosynthetic nitrogen-use efficiency was observed that could not be accounted for by changes in LMA. The gradual acclimation responses in the lower canopy may account, in whole or in part, for the approximately 2-year lag in post-harvest growth response observed in Acer saccharum.     

Tradeoff between shade adaptation and mitigation of photoinhibition in leaves of Quercus mongolica and Acer mono acclimated to deep shade

We investigated susceptibility to photoinhibition in leaves acclimated to different light regimes in intermediately shade-tolerant Japanese oak (Quercus mongolica Fisch. ex Turcz. var. crispula (Blume) Ohashi) and shade-tolerant Japanese maple (Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara), to elucidate adaptability to gap formation in leaves differing in shade acclimation. We hypothesized that there is a tradeoff between shade adaptation and capacity to mitigate photoinhibition associated with leaf morphology. We simultaneously measured chlorophyll fluorescence and gas exchange in seedlings that had been grown in full sunlight (open), 10% of full sun (moderate shade) and 5% of full sun (deep shade). Shade-tolerant A. mono adapted to deep shade through changes in leaf morphology, lowering its leaf mass per area (LMA), but Q. mongolica showed little change in LMA between moderate and deep shade. Photochemical quenching (qP) did not differ between species in full sunlight and moderate shade; however, in deep shade, qP of Q. mongolica was higher than that of A. mono, suggesting that Q. mongolica grown in deep shade is less susceptible to photoinhibition at gap formation. This is consistent with the finding that chronic photoinhibition 3 days after the transfer to full sunlight, indicated by the decrease in maximum photochemical efficiency, Fv/Fm, at predawn, was less in deep-shade-grown Q. mongolica than in deep shade-grown A. mono. In deep shade, the electron transport rate (ETR) of Q. mongolica was higher than that of A. mono, whereas thermal energy dissipation through photosystem II antennae, indicated by non-photochemical quenching, was lower in Q. mongolica than in A. mono. In deep shade, the greater ETR capacity in Q. mongolica in association with higher LMA and higher leaf N content could contribute to maintaining high qP and mitigating photoinhibition. These results indicate that, by maintaining a high electron transport capacity even in deep shade, the gap-dependent and intermediate-shade-tolerant Q. mongolica trades improved shade adaptation for higher growth potential when a gap event occurs.     

Leaf morphological and physiological adjustments to the spectrally selective shade imposed by anthocyanins in Prunus cerasifera

Prunus domestica L. has green leaves, whereas Prunus cerasifera Ehrh. var. atropurpurea has red leaves due to the presence of mesophyll anthocyanins. We compared morphological and photosynthetic characteristics of leaves of these species, which were sampled from shoots grafted in pairs on P. domestica rootstocks, each pair comprising one shoot of each species. Two hypotheses were tested: (1) anthocyanins protect red leaves against photoinhibition; and (2) red leaves display shade characteristics because of light attenuation by anthocyanins. Parameters were measured seasonally, during a period of increasing water stress, which caused a similar drop in shoot water potential in each species. As judged by predawn measurements of maximum PSII yield, chronic photoinhibition did not develop in either species and, despite the anthocyanic screen, the red leaves of P. cerasifera displayed lower light-adapted PSII yields and higher non-photochemical quenching than the green leaves of P. domestica. Thus, it appears that, in this system, anthocyanins afford little photoprotection. As predicted by the shade acclimation hypothesis, red leaves were thinner and had a lower stomatal frequency, area- based CO2 assimilation rate, apparent carboxylation efficiency and chlorophyll a:b ratio than green leaves. However, red leaves were similar to green leaves in conductivity to water vapor diffusion, dry-mass-based chlorophyll concentrations and carotenoid:chlorophyll ratios. The data for red leaves indicate adaptations to a green-depleted, red-enriched shade, rather than a neutral or canopy-like shade. Thus, green light attenuation by anthocyanins may impose a limitation on leaf thickness. Moreover, the selective depletion of light at wavelengths that are preferentially absorbed by PSII and chlorophyll b may lead to adjustments in chlorophyll pigment ratios to compensate for the uneven spectral distribution of internal light. The apparent photosynthetic cost associated with lost photons and reduced leaf thickness, and the absence of a photoprotective advantage, suggest that there are other, yet to be identified, benefits for permanently anthocyanic leaves of P. cerasifera.     

Leaf physiological versus morphological acclimation to high-light exposure at different stages of foliar development in oak

We investigated light acclimation in seedlings of the temperate oak Quercus petraea (Matt.) Liebl. and the co-occurring sub-Mediterranean oak Quercus pyrenaica Willd. Seedlings were raised in a greenhouse for 1 year in either 70 (HL) or 5.3% (LL) of ambient irradiance of full sunlight, and, in the following year, subsets of the LL-grown seedlings were transferred to HL either before leaf flushing (LL-HLBF plants) or after full leaf expansion (LL-HLAF plants). Gas exchange, chlorophyll a fluorescence, nitrogen fractions in photosynthetic components and leaf anatomy were examined in leaves of all seedlings 5 months after plants were moved from LL to HL. Differences between species in the acclimation of LL-grown plants to HL were minor. For LL-grown plants in HL, area-based photosynthetic capacity, maximum rate of carboxylation, maximum rate of electron transport and the effective photochemical quantum yield of photosystem II were comparable to those for plants grown solely in HL. A rapid change in nitrogen distribution among photosynthetic components was observed in LL-HLAF plants, which had the highest photosynthetic nitrogen-use efficiency. Increases in mesophyll thickness and dry mass per unit area governed leaf acclimation in LL-HLBF plants, which tended to have less nitrogen in photosynthetic components and a lower assimilation potential per unit of leaf mass or nitrogen than LL-HLAF plants. The data indicate that the phenological state of seedlings modified the acclimatory response of leaf attributes to increased irradiance. Morphological adaptation of leaves of LL-HLBF plants enhanced photosynthetic capacity per unit leaf area, but not per unit leaf dry mass, whereas substantial redistribution of nitrogen among photosynthetic components in leaves of LL-HLAF plants enhanced both mass- and area-based photosynthetic capacity.     

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.     

Sensitivity of photosynthetic electron transport to photoinhibition in a temperate deciduous forest canopy: Photosystem II center openness, non-radiative energy dissipation and excess irradiance under field conditions

We used chlorophyll fluorescence techniques to investigate responses of Photosystem II (PSII) quantum yield to light availability in the short term (quantum flux density integrated over the measurement day, Qd) and in the long term (Qd averaged over the season, Qs) in a mixed deciduous forest comprising shade-tolerant and water-stress-sensitive Tilia cordata Mill. in the lower canopy and shade-intolerant and water-stress-resistant Populus tremula L. in the upper canopy. In both species, intrinsic efficiency of PSII in the dark-adapted state (Fv/Fm) was lower during the day than during the night, and the difference in Fv/Fm between day and night increased with increasing Qs. Although the capacity for photosynthetic electron transport increased with increasing Qs in both species, maximum quantum efficiency of PSII in the light-adapted state (alpha) decreased with increasing Qs. At a common Qs, alpha was lower in T. cordata than in P. tremula primarily because of a higher fraction of closed PSII centers, and to a smaller extent because of limited, non-radiative, excitation energy dissipation in the pigment bed in T. cordata. Across both species, photochemical quenching (qP), which measures the openness of PSII centers, varied more than fivefold, but the efficiency of excitation energy capture by open PSII centers (Fv'/Fm'), which is an estimate of non-radiative excitation energy dissipation in PSII antennae, varied by only 50%. Chlorophyll turnover rates increased with increasing irradiance, especially in T. cordata, possibly because of increased photodestruction. Diurnal measurements of PSII quantum yields (PhiPSII) indicated that, under similar environmental conditions, PhiPSII was always lower in the afternoon than in the morning, and the fraction of daily integrated photosynthetic electron transport lost because of diurnal declines in PhiPSII (Delta) increased with increasing Qd. At a common Qd, mean daily PSII center reduction state, the fraction of light in excess (1 - fractions of light used in photochemistry and dissipated as heat) and Delta were higher in T. cordata than in P. tremula. This was attributed to greater stomatal closure during the day, which led to a greater reduction in the requirement for assimilative electron flow in T. cordata. Across both species, Delta scaled negatively with the fraction of light utilized photochemically, demonstrating the leading role of PSII center openness in maintaining high PSII efficiency. Because photosynthesis (A) at current ambient carbon dioxide concentration is limited by CO2 availability in high light and mainly by photosynthetic electron transport rates in low light, overall daily down-regulation of PhiPSII primarily influences A in low light. Given that foliar water stress scales positively with Qs in both species, we conclude that the inverse patterns of variation in water and light availabilities in the canopy result in a greater decline in A than is predicted by decreases in stomatal conductance alone.     

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.     

Interspecific variability of stem photosynthesis among tree species

The photosynthetic characteristics of current-year stems of six deciduous tree species, two evergreen tree species and ginkgo (Ginkgo biloba L.) were compared. Gas exchange, chlorophyll concentration, nitrogen concentration and maximum quantum yield of PSII were measured in stems in summer and winter. A light-induced decrease in stem CO(2) efflux was observed in all species. The apparent gross photosynthetic rate in saturating light ranged from 0.72 micromol m(-2 )s(-1) (ginkgo, in winter) to 3.73 micromol m(-2) s(-1) (Alnus glutinosa (L.) Gaertn., in summer). Despite this variability, a unique correlation (slope = 0.75), based on our results and those reported in the literature, was found between gross photosynthetic rate and dark respiration rate. Mass-based gross photosynthetic rate decreased with stem mass per area and correlated to chlorophyll concentration and nitrogen concentration, both in summer and winter. The radial distribution of stem chlorophyll differed among species, but all species except ginkgo had chlorophyll as deep as the pith. In summer, the maximum quantum yield of stem PSII (estimated from the ratio of variable to maximal fluorescence; F (v)/F (m)) of all species was near the optimal value found for leaves. By contrast, the values were highly variable in winter, suggesting large differences in sensitivity to low-temperature photoinhibition. The winter values of Fv/Fm were only 31-60% of summer values for the deciduous species, whereas the evergreen conifer species maintained high F (v)/F (m) in winter. The results highlight the interspecific variability of gross photosynthesis in the stem and its correlation with structural traits like those found for leaves. The structural correlations suggest that the selection of photosynthetic traits has operated under similar constraints in stems and leaves.     

Seasonal changes in photosynthesis and growth of Zizyphus attopensis seedlings in three contrasting microhabitats in a tropical seasonal rain forest

We hypothesized that photosynthesis and growth of tropical vegetation at its most northern distribution in Asia (Xishuangbanna, SW China) is adversely affected by seasonal drought and chilling temperatures. To test this hypothesis, we measured photosynthetic and growth characteristics of Zizyphus attopensis Pierre seedlings grown in three contrasting forest microhabitats: the understory, a small gap and a large gap. Photosynthetic capacity (light-saturated photosynthetic rate (A(max)), maximum rate of carboxylation and electron transport rate) and partitioning of leaf nitrogen (N) into carboxylation and electron transport differed significantly among seasons and microhabitats. Specific leaf area (SLA) did not change seasonally, but differed significantly among microhabitats and showed a negative linear relationship with daily integrated photon flux (PPF(i)). In contrast, leaf N concentration per unit area (N(a)) changed seasonally but did not differ among microhabitats. Measurements of maximum PSII photochemical efficiency (F(v)/F(m)) indicated that chronic photoinhibition did not occur in seedlings in any of the microhabitats during the study. Photosynthetic capacity was greatest in the wet season and lowest in the cool season. During the cool and dry seasons, the reduction in A(max) was greater in seedlings grown in the large gap than in in the understory and the small gap. Close logarithmic relationships were detected between PPF(i), leaf N(a) and photosynthetic capacity. Stem mass ratio decreased, and root mass ratio increased, in the dry season. We conclude that seasonal acclimation in growth and photosynthesis of the seedlings was associated with changes in biochemical features (particularly N(a) and partitioning of total leaf N between the different photosynthetic pools) and biomass allocation, rather than with changes in leaf morphological features (such as SLA). Local irradiance is the main factor driving seasonal variations in growth and photosynthesis in the study area, where the presence of heavy fog during the cool and dry seasons limits irradiance, but supplies water to the soil surface layers.     

Leaf morphology and photosynthetic adjustments among deciduous broad-leaved trees within the vertical canopy profile

Photosynthetic acclimation of deciduous broad-leaved tree species was studied along a vertical gradient within the canopy of a multi-species deciduous forest in northern Japan. We investigated variations in (1) local light regime and CO2 concentration ([CO2]), and (2) morphological (area, thickness and area per mass), biochemical (nitrogen and chlorophyll concentrations) and physiological (light-saturated photosynthetic rate) attributes of leaves of seven major species on three occasions (June, August and October). We studied early successional species, alder (Alnus hirsuta (Spach) Rupr.) and birch (Betula platyphylla var. japonica (Miq.) Hara); gap phase species, walnut (Juglans ailanthifolia Carrière) and ash (Fraxinus mandshurica var. japonica Rupr.); mid-successional species, basswood (Tilia japonica (Miq.) Simonk.) and elm (Ulmus davidiana var. japonica (Rehd.) Nakai); and the late-successional species, maple (Acer mono Bunge). All but maple initiated leaf unfolding from the lower part of the crown. The [CO2] within the vertical profile ranged from 320-350 ppm in the upper canopy to 405-560 ppm near the ground. The lowest and highest ambient [CO2] occurred during the day and during the night, respectively. This trend was observed consistently during the summer, but not when trees were leafless. Chlorophyll concentration was positively related to maximum photosynthetic rate within, but not among, species. Leaf senescence started from the inner part of the crown in alder and birch, but started either in the outer or top portion of the canopy of ash, basswood and maple. Chlorophyll (Chl) to nitrogen ratio in leaves increased with decreasing photon flux density. However, Chl b concentration in all species remained stable until the beginning of leaf senescence. Maximum photosynthetic rates observed in sun leaves of early successional species, gap phase or mid-successional species, and late successional species were 12.5-14.8 micromol m(-2) s(-1), 4.1-7.8 micromol m(-2) s(-1) and 3.1 micromol m(-2) s(-1), respectively.     

Three-dimensional lamina architecture alters light-harvesting efficiency in Fagus: a leaf-scale analysis

Modification of foliage exposition and morphology by seasonal average integrated quantum flux density (Qint) was investigated in the canopies of the shade-tolerant late-successional deciduous tree species Fagus orientalis Lipsky and Fagus sylvatica L. Because the leaves were not entirely flat anywhere in the canopy, the leaf lamina was considered to be three-dimensional and characterized by the cross-sectional angle between the leaf halves (theta). Both branch and lamina inclination angles with respect to the horizontal scaled positively with irradiance in the canopy, allowing light to penetrate to deeper canopy horizons. Lamina cross-sectional angle varied from 170 degrees in the most shaded leaves to 90-100 degrees in leaves in the top of the canopy. Thus, the degree of leaf rolling increased with increasing Qint, further reducing the light-interception efficiency of the upper-canopy leaves. Simulations of the dependence of foliage light-interception efficiency on theta demonstrated that decreases in theta primarily reduce the interception efficiency of direct irradiance, but that diffuse irradiance was equally efficiently intercepted over the entire range of theta values in our study. Despite strong alteration in foliage light-harvesting capacity within the canopy and greater transmittance of the upper crown compared with the lower canopy, mean incident irradiances varied more than 20-fold within the canopy, indicating inherent limitations in light partitioning within the canopy. This extensive canopy light gradient was paralleled by plastic changes in foliar structure and chemistry. Leaf dry mass per unit area varied 3-4-fold between the canopy top and bottom, providing an important means of scaling foliage nitrogen contents and photosynthetic capacity per unit area with Qint. Although leaf structure versus light relationships were qualitatively similar in all cases, there were important tree-to-tree and species-to-species variations, as well as evidence of differences in investments in structural compounds within the leaf lamina, possibly in response to contrasting leaf water availability in different trees.     

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.     

Energy investment in leaves of red maple and co-occurring oaks within a forested watershed

Despite its recent expansion in eastern US forests, red maple (Acer rubrum L.) generally exhibits a low leaf photosynthetic rate, leaf mass per unit area (LMA) and leaf nitrogen concentration ([N]) relative to co-occurring oaks (Quercus spp.). To evaluate these differences from the perspective of leaf energy investment, we compared leaf construction cost (CC) and leaf maintenance cost (MC) with leaf photosynthetic rate at saturating photon flux density and ambient CO2 partial pressure (Amax) in red maple and co-occurring red oak (Quercus rubra L.) and chestnut oak (Quercus prinus L.). We also examined relationships among leaf physiological, biochemical and structural characteristics of upper-canopy leaves of these three species at lower (wetter) and upper (drier) elevation sites of a watershed in the Black Rock Forest, Cornwall, NY, USA. Although A(max), leaf [N], leaf carbon concentration ([C]) and LMA were significantly less in red maple than in either oak species at both sites, CC per unit leaf area of red maple was 28.2 and 35.4% less than that of red oak at the lower and upper site, respectively, and 38.8 and 32% less than that of chestnut oak at the lower and upper site, respectively. Leaf MC per unit leaf area, which was positively associated with leaf CC (r2 = 0.95), was also significantly lower in red maple than in either oak species at both sites. When expressed per unit leaf area, A(max) was positively correlated with both CC (r2 = 0.65) and MC (r2 = 0.59). The cost/benefit ratio of CC/Amax of red maple was significantly less than that of chestnut oak at the lower site, however, CC/A(max) did not exhibit any significant interspecific differences at the upper site. Expressed per unit leaf area, CC was correlated positively with LMA (r2 = 0.90), leaf [N] (r2 = 0.97), and leaf [C] (r2 = 0.89), and negatively correlated with leaf molar carbon to nitrogen ratio (r2 = 0.92). Combined with red maple's general success in many oak-dominated forests, our findings suggest that reduced leaf-level photosynthetic capacity and related leaf characteristics in red maple are partially balanced by lower energy and resource requirements for leaf biomass construction and maintenance, which could enhance the competitive success of this species.     

Seasonal acclimation of photosystem II in Pinus sylvestris. I. Estimating the rate constants of sustained thermal energy dissipation and photochemistry

Acclimation of the partitioning of absorbed light energy in Photosystem II (PSII) between photochemical and non-photochemical processes includes short-term adjustments that are rapidly reversed in the dark and seasonal acclimation processes that are unaffected by dark acclimation. Thus, by using dark-acclimated leaves to study the seasonal acclimation of PSII, the confounding effect of short-term adjustments is eliminated. The maximum quantum yield of photochemistry, estimated by chlorophyll fluorescence analysis as F(v)/F(m), where F(v) = (F(m) - F(o)), and F(m) and F(o) are maximum and minimum chlorophyll fluorescence, respectively, has been widely used to follow the seasonal acclimation of PSII, because it is measured in dark-acclimated leaves. Seasonal changes in F(v)/F(m) can be caused by adjustments in either the photochemical capacity in PSII, or the capacity of thermal dissipation in PSII, or both. However, there is a lack of chlorophyll fluorescence parameters that can distinguish between these processes. In this study, we introduce two new parameters: the rate constants of sustained thermal energy dissipation (k(NPQ)) and of photochemistry (k(P)). We estimated k(NPQ) and k(P) from dark-acclimated F(o) and F(m) measured during spring recovery of photosynthesis in Scots pine (Pinus sylvestris L.) trees. We suggest that k(NPQ) and k(P) be used to study the mechanisms underlying the observed seasonal acclimation in PSII, because these parameters provide quantitative data that complement and extend F(v)/F(m) measurements.     

Photosynthetic acclimation of overstory Populus tremuloides and understory Acer saccharum to elevated atmospheric CO2 concentration: interactions with shade and soil nitrogen

We exposed Populus tremuloides Michx. and Acer saccharum Marsh. to a factorial combination of ambient and elevated atmospheric CO2 concentrations ([CO2]) and high-nitrogen (N) and low-N soil treatments in open-top chambers for 3 years. Our objective was to compare photosynthetic acclimation to elevated [CO2] between species of contrasting shade tolerance, and to determine if soil N or shading modify the acclimation response. Sun and shade leaf responses to elevated [CO2] and soil N were compared between upper and lower canopy leaves of P. tremuloides and between A. saccharum seedlings grown with and without shading by P. tremuloides. Both species had higher leaf N concentrations and photosynthetic rates in high-N soil than in low-N soil, and these characteristics were higher for P. tremuloides than for A. saccharum. Electron transport capacity (Jmax) and carboxylation capacity (Vcmax) generally decreased with atmospheric CO2 enrichment in all 3 years of the experiment, but there was no evidence that elevated [CO2] altered the relationship between them. On a leaf area basis, both Jmax and Vcmax acclimated to elevated [CO2] more strongly in shade leaves than in sun leaves of P. tremuloides. However, the apparent [CO2] x shade interaction was largely driven by differences in specific leaf area (m2 g-1) between sun and shade leaves. In A. saccharum, photosynthesis acclimated more strongly to elevated [CO2] in sun leaves than in shade leaves on both leaf area and mass bases. We conclude that trees rooted freely in the ground can exhibit photosynthetic acclimation to elevated [CO2], and the response may be modified by light environment. The hypothesis that photosynthesis acclimates more completely to elevated [CO2] in shade-tolerant species than in shade-intolerant species was not supported.     

Leaf phenology,photosynthesis, and the persistence of saplings and shrubs in a mature northern hardwood forest

We quantified leaf phenologies of saplings and overstory trees of sugar maple (Acer saccharum Marsh.) and American beech (Fagus grandifolia Ehrh.), and the shrub hobblebush viburnum (Viburnum alnifolium Marsh.) in a 72-year-old northern hardwood forest. Seasonal changes in irradiance in the shrub layer, and in the leaf CO(2) exchange of viburnum, and sugar maple and beech saplings were also measured. Leaf expansion occurred earlier in the spring and green leaves were retained later in the autumn in saplings and shrubs than in overstory trees. During the spring light phase (before overstory closure), large CO(2) gains by all three shrub-layer species occurred as a result of a combination of relatively large leaf area, high photosynthetic capacity, and high irradiance. Throughout the summer shade phase, photosynthetic capacity at a given irradiance remained relatively constant, but CO(2) gain was typically limited by low irradiances. Even though irradiance in the shrub layer increased during the autumn light phase as the overstory opened, CO(2) gains were modest compared to springtime values because of declining leaf area and photosynthetic capacity in all three species. The CO(2) gains during the spring light phase, and to a lesser extent during the autumn light phase, may be important to the carbon balance and long-term persistence of saplings and shrubs in the usually light-limited shrub layer of a northern hardwood forest. Therefore, for some late-successional species, leaf phenology may be an important characteristic that permits their long-term persistence in the shrub layer of mature northern hardwood forests.     

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.