Effect of thinning on anatomical adaptations of Norway spruce needles

Conifers and other trees are constantly adapting to changes in light conditions, water/nutrient supply and temperatures by physiological and morphological modifications of their foliage. However, the relationship between physiological processes and anatomical characteristics of foliage has been little explored in trees. In this study we evaluated needle structure and function in Norway spruce families exposed to different light conditions and transpiration regimes. We compared needle characteristics of sun-exposed and shaded current-year needles in a control plot and a thinned plot with 50% reduction in stand density. Whole-tree transpiration rates remained similar across plots, but increased transpiration of lower branches after thinning implies that sun-exposed needles in the thinned plot were subjected to higher water stress than sun-exposed needles in the control plot. In general, morphological and anatomical needle parameters increased with increasing tree height and light intensity. Needle width, needle cross-section area, needle stele area and needle flatness (the ratio of needle thickness to needle width) differed most between the upper and lower canopy. The parameters that were most sensitive to the altered needle water status of the upper canopy after thinning were needle thickness, needle flatness and percentage of stele area in needle area. These results show that studies comparing needle structure or function between tree species should consider not only tree height and light gradients, but also needle water status. Unaccounted for differences in needle water status may have contributed to the variable relationship between needle structure and irradiance that has been observed among conifers.     

Acclimation of Pacific yew (Taxus brevifolia) foliage to sun and shade

The success in clinical trials of the anti-cancer drug, Taxol(R), obtained from the bark of Pacific yew (Taxus brevifolia Nutt.), has raised interest in cultivation and regeneration of this little-known species. Pacific yew is shade-tolerant and it is not known whether the foliage can tolerate the high solar irradiances found on an open forest regeneration site or a nursery. Acclimation of Pacific yew to sun and shade was studied by comparing foliar physiology and morphology of male and female trees growing in full sun or shade. Interspecific foliar acclimation to sun was studied by comparing sun-grown English yew (Taxus baccata L.) with Pacific yew. No sex-specific acclimation was found in foliar physiology or morphology in either species. Sun-grown foliage of Pacific yew and English yew differed with respect to light harvesting, transpiration, stomatal conductance, leaf structure, stomatal distribution and foliar N concentrations and contents. Chlorophyll a fluorescence measurements indicated that shade-grown foliage of Pacific yew had larger and more efficient light harvesting systems than sun-grown foliage. Rates of CO(2) uptake and transpiration were similar in sun- and shade-grown foliage indicating acclimation of photosynthesis to the growth irradiance. Specific leaf area was significantly higher in shade-grown foliage of Pacific yew than in sun-grown foliage and was diagnostic of the light environment in which the foliage grew. Foliar N concentrations were not significantly different between sun- and shade-grown leaves of Pacific yew but sun-grown foliage had a higher N content. Physiological and morphological adjustments of Pacific yew foliage conferred tolerance to both high light and shade, enabling the trees to survive in a variety of light environments and indicating that Pacific yew is suited to nursery cultivation and regeneration of open sites.     

Rapid freezing injury in red spruce: seasonal changes in sensitivity and effects of temperature range

On calm, cold days in winter, sun-exposed needles of red spruce (Picea rubens Sarg.) may warm 10 to 20 degrees C above ambient air temperature, and undergo rapid (>/= 1 degrees C min(-1)) fluctuations in temperature as light breezes or passing clouds alter the energy balance of the foliage. It has been proposed that the resulting rapid freeze-thaw cycles (freezing stress) cause a type of winter injury in montane red spruce that is characterized by necrosis of sun-exposed foliage. In autumn and winter, we monitored rapid freezing stress response of needle sections from 10 montane red spruce trees by subjecting needles to rapid freezing over the temperature span typically recorded in the field. In autumn, experimental rapid freezing stress produced severe injury only at temperatures considerably lower than expected for that time of year. In winter, rapid freezing caused occasional, moderate injury in fully hardened foliage of trees susceptible to both slow and rapid freezing. Seasonal changes in sensitivity to rapid and slow freezing were correlated, suggesting that environmental factors that are known to affect sensitivity to slow freezing may also affect sensitivity to rapid freezing. Experimental manipulation of the start and end temperatures of rapid freezing stress events showed that moderate to severe needle injury can occur in susceptible trees at temperature spans slightly more extreme than those typically recorded in the field. The extent of injury was similar at different starting temperatures if rapid freezing occurred over the same temperature span. Year-old foliage was consistently less sensitive to rapid freezing stress than current-year foliage, but some year-old foliage was damaged when the rapid freezing stress regime caused severe injury in current-year foliage. We conclude that rapid freeze-thaw cycles can explain light to moderate injury of current-year foliage, but they do not explain the more severe and widespread pattern of foliar damage that has occurred intermittently over at least the last 18 years.     

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.     

Dependence of needle architecture and chemical composition on canopy light availability in three North American Pinus species with contrasting needle length

Morphology and chemical composition of needles of shade-intolerant southern conifers (Pinus palustris Mill. (mean needle length +/- SD = 29.1 +/- 4.1 cm), P. taeda L. (12.3 +/- 2.9 cm) and P. virginiana Mill. (5.1 +/- 0.8 cm)) were studied to test the hypothesis that foliage acclimation potential to canopy light gradients is generally low for shade-intolerant species, and in particular, because of mechanical limitations, in species with longer needles. Plasticity for each needle variable was defined as the slope of the foliar characteristic versus irradiance relationship. A novel geometrical model for needle area and volume calculation was employed for the three-needled species P. palustris and P. taeda. Needle thickness (T) strongly increased, but width (W) was less variable with increasing daily integrated quantum flux density averaged over the season (Q(int)), resulting in changes in cross-sectional needle shape that were manifested in a positive relationship between the total to projected needle area ratio (A(T)/A(P)) and Q(int) in the three-needled species. In contrast, cross-sectional needle geometry was only slightly modified by irradiance in the two-needled conifer P. virginiana. Needle dry mass per unit total needle area (M(T)) was positively related to Q(int) in all species, leading to greater foliar nitrogen contents per unit area at higher irradiances. Separate examination of the components of M(T) (density (D) and the volume (V) to A(T) ratio; M(T) = DV/A(T)) indicated that the positive effect of light on M(T) resulted solely from increases in V/A(T), i.e., from increases in the thickness of foliage elements. Foliar chlorophyll content per unit mass increased with increasing Q(int), allowing an improvement in light-harvesting efficiency in low light. The variables characterizing needle material properties (D, the dry to fresh mass ratio, and needle carbon content per unit mass) were generally independent of Q(int), suggesting that needles were less stiff and had greater tip deflections under their own weight at lower irradiances because of smaller W and T. Comparisons with the literature revealed that plasticity in foliar characteristics tended to be lower in the studied shade- intolerant species than in shade-tolerant conifers, but plasticity among the investigated species was unaffected by needle length. However, we argue that, because of mechanical limitations, plastic changes in needle cross section in response to low irradiance may decrease rather than increase light-interception efficiency in long-needled species.     

Stomatal conductance alone does not explain the decline in foliar photosynthetic rates with increasing tree age and size in Picea abies and Pinus sylvestris

Foliar light-saturated net assimilation rates (A) generally decrease with increasing tree height (H) and tree age (Y), but it is unclear whether the decline in A is attributable to size- and age-related modifications in foliage morphology (needle dry mass per unit projected area; M(A)), nitrogen concentration, stomatal conductance to water vapor (G), or biochemical foliage potentials for photosynthesis (maximum carboxylase activity of Rubisco; V(cmax)). I studied the influences of H and Y on foliage structure and function in a data set consisting of 114 published studies reporting observations on more than 200 specimens of various height and age of Picea abies (L.) Karst. and Pinus sylvestris L. In this data set, foliar nitrogen concentrations were independent of H and Y, but net assimilation rates per unit needle dry mass (A(M)) decreased strongly with increasing H and Y. Although M(A) scaled positively with H and Y, net assimilation rates per unit area (A(A) = M(A) x A(M)) were strongly and negatively related to H, indicating that the structural adjustment of needles did not compensate for the decline in mass-based needle photosynthetic rates. A relevant determinant of tree height- and age-dependent modifications of A was the decrease in G. This led to lower needle intercellular CO2 concentrations and thereby to lower efficiency with which the biochemical photosynthetic apparatus functioned. However, V(cmax) per unit needle dry mass and area strongly decreased with increasing H, indicating that foliar photosynthetic potentials were lower in larger trees at a common intercellular CO2 concentration. Given the constancy of foliar nitrogen concentrations, but the large decline in apparent V(cmax) with tree size and age, I hypothesize that the decline in Vcmax results from increasing diffusive resistances between the needle intercellular air space and carboxylation sites in chloroplasts. Increased diffusive limitations may be the inevitable consequence of morphological adaptation (changes in M(A) and needle density) to greater water stress in needles of larger trees. Foliage structural and physiological variables were nonlinearly related to H and Y, possibly because of hyperbolic decreases in shoot hydraulic conductances with increasing tree height and age. Although H and Y were correlated, foliar characteristics were generally more strongly related to H than to Y, suggesting that increases in height rather than age are responsible for declines in foliar net assimilation capacities.     

Influence of nutrient supply on shade-sun acclimation of Picea abies seedlings: effects on foliar morphology,photosynthetic performance and growth

Norway spruce seedlings (Picea abies Karst.) were grown in low light for one year, under conditions of adequate and limiting nutrition, then transferred to high light. Three months after transfer we measured photosynthesis, leaf nitrogen concentration, leaf chlorophyll concentration and leaf mass per area (LMA) of current-year and 1-year-old shoots; silhouette area ratio (SAR, the ratio of shoot silhouette area to projected needle area) was also measured in current-year shoots. At the foliage level, the effects of light and nutrient treatments differed markedly. Light availability during foliage expansion primarily affected LMA and SAR (morphological acclimation at the needle and shoot level, respectively). By contrast, nutrient supply in high light affected photosynthetic capacity per unit of leaf tissue (physiological acclimation at the cellular level) but did not affect LMA and SAR. The capacity for shade-sun acclimation in foliage formed before transfer to high light differed greatly from that of foliage formed following the transfer. The morphological inflexibility of mature needles (measured by LMA) limited their shade-sun acclimation potential. In contrast, at high nutrient supply, shoots that developed just after the change in photosynthetic photon flux density largely acclimated, both morphologically and physiologically, to the new light environment. The acclimation response of both current- and 1-year-old shoots was prevented by nutrient limitation. Analysis of growth at the whole-plant level largely confirmed the conclusions drawn at the shoot level. We conclude that nutrient shortage subsequent to the opening of a canopy gap may strongly limit the acclimation response of Norway spruce seedlings. Successful acclimation was largely related to the plant's ability to produce sun foliage and adjust whole-plant biomass allocation rapidly.     

Effects of elevated CO(2) concentration and nutrition on net photosynthesis,stomatal conductance and needle respiration of field-grown Norway spruce trees

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.     

Plasticity of shoot and needle morphology and photosynthesis of two Picea species with different site preferences in northern Japan

We compared shoot and needle morphology and photosynthesis in Picea glehnii (Friedr. Schmidt) M.T. Mast. and Picea jezoensis (Sieb. et Zucc.) Carr. trees planted on immature volcanic ash and well-developed brown forest soils to investigate whether species differences in morphological and physiological plasticity affected tree growth on different soil types. Height growth of P. glehnii was reduced by about 10% on volcanic ash compared with forest soil, whereas that of P. jezoensis was reduced by more than 60%. Needle morphology of P. glehnii was unaffected by soil type. In contrast, needles of P. jezoensis trees growing on volcanic ash were shorter, narrower and thicker, with less dry mass and area, than those of trees growing on forest soil, and specific needle area was lower, indicating lower foliar light-interception efficiency. In both species, changes in needle morphology with increasing irradiance were similar in trees growing on both soil types, indicating that plasticity of needle morphology was unaffected by soil type. In both species, shoot mass and shoot silhouette area were lower and needle mass per unit shoot mass was higher in trees growing on volcanic ash than in trees growing on forest soil. Trees of both species had more needles per unit shoot length, lower shoot silhouette to projected needle area ratios and lower shoot silhouette areas per unit shoot mass (SAM) on volcanic ash than on forest soil, indicating lower shoot-level light-interception efficiency. For P. glehnii, the response of shoot morphology to increasing irradiance was similar on both soil types, with the exception of SAM, which showed lower plasticity in trees growing on volcanic ash. In contrast, shoot-level morphological plasticity of P. jezoensis was reduced in trees growing on volcanic ash. Light-saturated maximum photosynthetic rate (P(max)) of P. glehnii was unaffected by soil type, whereas mass-based P(max) of P. jezoensis was lower in trees growing on volcanic ash than in trees growing on forest soil. In P. jezoensis trees growing on forest soil, area-based P(max) increased with increasing irradiance, but this response was not observed in trees growing on volcanic ash. As a result, area-based P(max) at the top of the canopy was 39 to 54% lower in trees growing on volcanic ash than in trees growing on forest soil. Our results indicate that constraints on morphological acclimation to high irradiances may contribute to reduced height growth of P. jezoensis on volcanic ash.     

Seasonal acclimation of photosystem II in Pinus sylvestris. II. Using the rate constants of sustained thermal energy dissipation and photochemistry to study the effect of the light environment

Photosynthesis in evergreen conifers is characterized by down-regulation in autumn and rapid up-regulation in spring. This seasonal pattern is largely driven by temperature, but the light environment also plays a role. In overwintering Scots pine (Pinus sylvestris L.) trees, PSII is less down-regulated and recovers faster from winter stress in shaded needles than in needles exposed to full sunlight. Because the effect of light on the seasonal acclimation of PSII has not been quantitatively studied under field conditions, we used the rate constants for sustained thermal energy dissipation and photochemistry to investigate the dynamics and kinetics of the seasonal acclimation of PSII in needles exposed to different light environments. We monitored chlorophyll fluorescence and needle pigment concentration during the winter and spring in Scots pine seedlings growing in the field in different shading treatments, and within the crowns of mature trees. The results indicated that differences in acclimation of PSII in overwintering Scots pine among needles exposed to different light environments can be chiefly attributed to sustained thermal dissipation. We also present field evidence that zeaxanthin-facilitated thermal dissipation and aggregation of thylakoid membrane proteins are key mechanisms in the regulation of sustained thermal dissipation in Scots pine trees in the field.     

Effects of foliar potassium concentration on morphology, ultrastructure and polyamine concentrations of Scots pine needles

We examined the effects of three foliar potassium concentrations (high, intermediate and low) on the morphology, ultrastructure and polyamine concentrations of current-year and 1- and 2-year-old needles of 30-year-old Scots pine (Pinus sylvestris L.) trees. Foliar K concentration had only a slight effect on needle morphology. The sclerenchyma cell walls were thinner, the xylem area was larger, and the resin ducts were smaller in needles with a low K concentration than in needles with a high or intermediate K concentration. In addition, the bundle sheath cells were collapsed in needles having a low K concentration. The secondary growth of phloem tissue and the mesophyll area were greater in needles with a high or intermediate K concentration than in needles with a low K concentration, possibly indicating greater production of photoassimilates in these trees. At the ultrastructural level, mesophyll cells with enlarged central vacuoles and small vacuoles containing electron-dense material were common in needles having a low K concentration. Enlargement of the central vacuole coincided with an exponential increase in putrescine concentration in needles with a low K concentration, suggesting that the central vacuole may function as a storage site for putrescine.     

The effects of ultraviolet-B radiation on photosynthesis of different aged needles in field-grown loblolly pine

We examined the effect of supplemental UV-B radiation (290-320 nm) on photosynthetic characteristics of different aged needles of 3-year-old, field-grown loblolly pine (Pinus taeda L.). Needles in four age classes were examined: I, most recently fully expanded, year 3; II, first flush, year 3; III, final flush, year 2; and IV, oldest needles still present, year 2. Enhanced UV-B radiation caused a statistically significant decrease (6%) in the ratio of variable to maximum fluorescence (F(v)/F(m)) following dark adaptation only in needles from the youngest age class, suggesting transient damage to photosynthesis. However, no effects of enhanced UV-B radiation on other instantaneous measures of photosynthesis, including maximum photosynthesis, apparent quantum yield and dark respiration, were seen for needles of any age. Foliar nitrogen concentration was unaffected by UV-B treatment. However, the (13)C/(12)C carbon isotope ratios (delta(13)C-a time integrated measure of photosynthetic function) of needles in age classes II and IV were 3% (P < 0.01) and 2% (P < 0.05) more negative, respectively, in treated plants than in control plants. Exposure to enhanced UV-B radiation caused a 20% decrease in total biomass and a 4% (P < 0.05), 25% (P < 0.01), and 9% (P < 0.01) decrease in needle length of needles in age classes I, II, and IV, respectively. The observed decreases in delta(13)C, and F(v)/F(m) of the needles in the youngest needle age class suggest subtle damage to photosynthesis, although overall growth reductions were probably a result of decreased total leaf surface rather than decreased photosynthetic capacity. Needles of age class IV had lower light- and CO(2)-saturated maximum photosynthetic rates (39%), lower dark respiration (34%), lower light saturation points (37%), lower foliar nitrogen concentration (28%), and lower delta(13)C (14%) values than needles of age class I. Apparent quantum yield and F(v)/F(m) did not change with needle age. The observed changes in photosynthesis and foliage chemical composition with needle age are consistent with previous studies of coniferous trees and may represent adaptations of older needles to shaded conditions within the canopy.     

Nutrient contents and concentrations in relation to growth of Picea abies and Fagus sylvatica along a European transect

Mineral nutrition of Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) was investigated along a transect extending from northern Sweden to central Italy. Nitrogen (N) concentrations of needles and leaves in stands growing on acid soils did not differ significantly between central Italy and southern Sweden (1.0 +/- 0.1 mmol N g(-1) for needles and 1.9 +/- 0.14 mmol N g(-1) for leaves). In both species, foliar N concentrations were highest in Germany (1.2 mmol N g(-1) for needles and 2.0 mmol N g(-1) for leaves) and decreased by 50% toward northern Sweden (0.5 mmol N g(-1)). Both species showed constant S/N and P/N ratios along the transect. Calcium, K and Mg concentrations generally reflected local soil conditions; however, Mg concentrations reached deficiency values in Germany. Leaf area per unit dry weight varied significantly along the transect with lowest values for Norway spruce recorded in northern Sweden and Italy (3.4 m(2) kg(-1)) and a maximum in central Europe (4.7 m(2) kg(-1)). A similar pattern was observed for beech. Despite the low variation in foliar N concentrations on the large geographic scale, local and regional variations in N concentrations equalled or exceeded the variation along the entire continental transect. Furthermore, nutrient contents (i.e., nutrient concentration x dry weight per needle or leaf) showed a greater variation than nutrient concentrations along the transect. Nitrogen contents of Norway spruce needles reached minimum values in northern Sweden (2.4 micro mol N needle(-1)) and maximum values in Denmark (5.0 micro mol N needle(-1)). The N content of beech leaves was highest in Denmark (242 micro mol N leaf(-1)). At the German site, foliar N content rather than N concentration reflected the seasonal dynamics of foliar growth and N storage of the two species. During foliage expansion, there was an initial rapid increase in N content and a decrease in N concentration. This pattern lasted for about 2 weeks after bud break and was followed by 6 weeks during which dry weight and N content of the foliage increased, resulting in a further decrease in N concentration. During summer, dry weight and N content of mature needles of Norway spruce increased further to reach a maximum in autumn, whereas N concentration remained constant. In spring, reallocation of N from 1- and 2-year-old needles was 1.5 and 1.0 micro mol N needle(-1), respectively. This remobilized N was a major source of N for the development of new needles, which had an N content of 1.5 micro mol N needle(-1) after bud break. The seasonal remobilization of N from old foliage decreased with increasing needle age. Needle N content and dry weight decreased progressively with age (1 micro mol N needle(-1) between age classes 2 and 5), whereas N concentrations remained constant. For Norway spruce, annual stemwood production was correlated with needle N content but not with foliar N concentration or with the total amount of N in the canopy. Interspecific and geographical differences in plant nutrition are discussed on the basis of competitive demands for C and N between growth of foliage and wood.     

Shoot structure,light interception,and distribution of nitrogen in an Abies amabilis canopy

We studied the effects of variation in shoot structure and needle morphology on the distributions of light and nitrogen within a Pacific silver fir (Abies amabilis (Dougl.) Forbes) canopy. Specifically, we investigated the role of morphological shade acclimation in the determination of resource use efficiency, which is claimed to be optimal when the distribution of nitrogen within the canopy is directly proportional to the distribution of intercepted photosynthetically active radiation (PAR). Shoots were collected from different heights in the crowns of trees representing four different size classes. A new method was developed to estimate seasonal light interceptance (SLI, intercepted PAR per unit needle area) of the shoots using a model for the directional distribution of above-canopy PAR, measurements of shoot silhouette area and canopy gap fraction in different directions. The ratio SLI/SLI(o), where the reference value SLI(o) represents the seasonal light interceptance of a spherical surface at the shoot location, was used to quantify the efficiency of light capture by a shoot. The ratio SLI/SLI(o) doubled from the top to the bottom of the canopy, mainly as a result of smaller internal shading in shade shoots than in sun shoots. Increased light-capturing efficiency of shade shoots implies that the difference in intercepted light by sun shoots versus shade shoots is much less than the decrease in available light from the upper to the lower canopy. For example, SLI of the five most sunlit shoots was only about 20 times greater than the SLI of the five most shaded shoots, whereas SLI(o) was 40 times greater for sun shoots than for shade shoots. Nitrogen content per unit needle area was about three times higher in sun needles than in shade needles. This variation, however, was not enough to produce proportionality between the amounts of nitrogen and intercepted PAR throughout the canopy.     

Canopy position and needle age affect photosynthetic response in field-grown Pinus radiata after five years of exposure to elevated carbon dioxide partial pressure

Photosynthesis of tree seedlings is generally enhanced during short-term exposure to elevated atmospheric CO2 partial pressure, but longer-term studies often indicate some degree of photosynthetic adjustment. We present physiological and biochemical evidence to explain observed long-term photosynthetic responses to elevated CO2 partial pressure as influenced by needle age and canopy position. We grew Pinus radiata D. Don. trees in open-top chambers for 5 years in sandy soil at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressures. The trees were well watered and exposed to natural light and ambient temperature. In the fourth year of CO2 exposure (fall 1997), when foliage growth had ceased for the year, photosynthetic down-regulation was observed in 1-year-old needles, but not in current-year needles, suggesting a reduction in carbohydrate sink strength as a result of increasing needle age (Turnbull et al. 1998). In 5-year-old trees (spring 1997), when foliage expansion was occurring, photosynthetic down-regulation was not observed, reflecting significantly large sinks for carbohydrates throughout the tree. Net photosynthesis was stimulated by 79% in trees growing in elevated CO2 partial pressure, but there was no significant effect on photosynthetic capacity or Rubisco activity and concentration. Current-year needles were more responsive to elevated CO2 partial pressure than 1-year-old needles, exhibiting larger relative increases in net photosynthesis to elevated CO2 partial pressure (98 versus 64%). Lower canopy and upper canopy leaves exhibited similar relative responses to growth in elevated CO2 partial pressure. However, needles in the upper canopy exhibited higher net photosynthesis, photosynthetic capacity, and Rubisco activity and concentration than needles in the lower canopy. Given that the ratio of mature to juvenile foliage mass in the canopy will increase as trees mature, we suggest that trees may become less responsive to elevated CO2 partial pressure with increasing age. We conclude that tree response to elevated CO2 partial pressure is based primarily on sink strength and not on the duration of exposure.     

Age-related changes in foliar morphology and physiology in red spruce and their influence on declining photosynthetic rates and productivity with tree age

The contribution of changes in meristem behavior to age-related decline in forest productivity is poorly understood. We studied age-related trends in needle morphology and gas exchange in a population of red spruce (Picea rubens Sarg.) growing in a multi-cohort stand where trees ranged from first-year germinants to trees over 150 years old, as well as in grafted scions from these trees. In the field study, age-related trends in foliar morphology were determined in six cohorts ranging in age from 2 to 120 years, and differences in gas exchange characteristics were compared between 60- and 120-year age classes. In a common-rootstock study, scions from trees representing 20-, 60-, and 120-year cohorts were grafted onto juvenile rootstock and maintained for three growing seasons, after which morphological and physiological foliar attributes were evaluated. The field study revealed significant age-related trends in foliar morphology, including decreasing specific leaf area, and increasing needle width, projected area, and width/length ratio. Similar trends were apparent in foliage from the grafted scions. Both in situ foliage and shoots of grafted scions from the oldest cohort showed significantly lower photosynthetic rates than their counterparts from younger trees; however, differences in stomatal conductance and internal CO(2) concentrations were not significant. These results suggest that: (1) foliage of red spruce exhibits age-related trends in both morphology and physiology; (2) age-related decreases in photosynthetic rates contribute to declining productivity in old red spruce; (3) declines in photosynthetic rates result from nonstomatal limitations; and (4) age-related changes in morphology and physiology are inherent in meristems and persist for at least 3 years in scions grafted to juvenile rootstock.     

Foliar morphological and physiological plasticity in Picea abies and Abies alba saplings along a natural light gradient

The role of morphological versus physiological foliar plasticity in the capacity for, and mechanisms of, photosynthetic acclimation was assessed in Picea abies (L.) Karst. and Abies alba Mill. saplings in a forest gap-understory light gradient (relative irradiance, RI, ranging from 0.02 to 0.32). The species investigated showed a similar foliar morphological plasticity along the light gradient, at both the needle level (through alteration in leaf dry mass per area) and the shoot level (through alteration in the silhouette area ratio, e.g., shoot silhouette to projected needle area ratio). In both species chlorophyll (Chl) concentration on a mass basis decreased at increasing RI, but was independent of RI when expressed on an area basis. In contrast, leaf N concentration on a mass basis was independent of RI, but was positively influenced by RI when expressed on an area basis. The parameters describing photosynthetic performance at low light (dark respiration rate, apparent quantum yield and light compensation point) suggest that Abies alba was better suited to maintain a positive carbon balance in shaded conditions. By contrast, parameters describing biochemical capacity at high light (maximum electron transport rate, Jmax and maximum ribulose-1,5-biphosphate carboxylation capacity, Vcmax) indicate that only Picea abies was capable of acclimating physiologically to high photosynthetic photon flux densities (PPFDs) by increasing nitrogen partitioning to Rubisco and Vcmax/mass by increasing RI. These results support the hypothesis that interspecific differences in nitrogen partitioning within the photosynthetic apparatus may provide a mechanistic basis for species separation along a light gradient. The differences in photosynthetic plasticity observed are likely to influence regeneration patterns and habitat breadth of the species investigated. The limited ability of Abies alba saplings to exploit high-light conditions may be a competitive disadvantage in large canopy gaps and thus limit recruitment of this species to small gaps.     

Acclimation of shoot and needle morphology and photosynthesis of two Picea species to differences in soil nutrient availability

To investigate morphological acclimation to differences in nutrient availability, we compared shoot and needle morphology of Picea glehnii (Friedr. Schmidt) M. T. Mast. and Picea jezoensis (Siebold & Zucc.) Carrière trees growing on nutrient-poor volcanic ash and nutrient-rich, brown forest soil. Trees of both species were shorter and had more open canopies when growing on volcanic ash than when growing on brown forest soil. Nutrient-poor conditions limited height growth less in P. glehnii than in P. jezoensis. In both species, trees growing on volcanic ash had shorter annual increments in the previous year and more needles per shoot length and, hence, a smaller shoot silhouette area (SSA) relative to needle dry mass (NDM) than trees growing on brown forest soil. Soil type had less effect on shoot projected needle area (PNA). Total needle area (TNA) of P. glehnii shoots was similar between soil types, whereas TNA of P. jezoensis was lower in trees growing on volcanic ash than in trees growing on brown forest soil. For both species, low SSA in response to nutrient-poor conditions resulted in low shoot SSA/PNA ratios, indicating high within-shoot self-shading. Shoot SSA/TNA of P. glehnii was lower in trees growing on volcanic ash than in trees growing on brown forest soil, indicating that needles were sun-acclimated. In contrast, shoot SSA/TNA of P. jezoensis was higher in trees growing on volcanic ash than in trees growing on brown forest soil. The contrasting response of TNA to low nutrient availability was associated with species-specific differences in needle morphology. Needles of P. glehnii growing on volcanic ash were slightly shorter, wider, thicker and heavier than those of trees growing on brown forest soil, indicating morphological acclimation to high irradiance. Needles of P. jezoensis growing on volcanic ash were shorter than those of trees growing on brown forest soil, but did not show morphological acclimation to high irradiance in width, thickness or mass. For both species, nutrient-poor conditions decreased maximum photosynthetic rate (Amax) per NDM. However, when expressed per PNA, the decrease in Amax was reduced, and when expressed per SSA, Amax was higher in trees growing on volcanic ash than in trees growing on brown forest soil. On volcanic ash, Amax per NDM was lower for P. glehnii than for P. jezoensis. However, morphological changes at the shoot and needle levels reversed this trend when Amax was expressed per SSA or per PNA. The species-specific differences in morphological response to differences in soil nutrient availability suggest that P. glehnii is more tolerant of nutrient-poor conditions, whereas P. jezoensis is better at exploiting nutrient-rich soils.     

Variability of the photosynthetic characteristics of mature needles within the crown of a 25-year-old Pinus pinaster

Photosynthetic characteristics of 1- and 2-year-old needles were determined in excised shoots of maritime pine (Pinus pinaster Ait.) with an open gas exchange system. We used the nonlinear least mean squares method to derive values for quantum yield of electron transport (alpha), maximum carboxylation velocity (V(cmax)), and maximum electron transport rate (J(max)), from photosynthetic response curves to light and CO(2). Crown height had no significant effect on any of the parameters; however, V(cmax) and J(max), as well as alpha were 43, 26 and 35% higher, respectively, in 1-year-old needles than in 2-year-old needles. The main effect of irradiance on needles was a small decline in leaf concentrations of nitrogen and phosphorus from the top to the bottom of the canopy. Only J(max) demonstrated a linear relationship with both nitrogen content (R(2) = 0.42) and irradiance at the shoot level. Because needle age accounted for most of the variability in photosynthesis, we incorporated needle age into the photosynthesis model of Farquhar et al. (1980). The modified model underestimated the daily assimilation rate of 1-year-old needles in the field, especially when assimilation rates were high.     

An analysis of light effects on foliar morphology,physiology, and light interception in temperate deciduous woody species of contrasting shade tolerance

Maximum Rubisco activities (V(cmax)), rates of photosynthetic electron transport (J(max)), and leaf nitrogen and chlorophyll concentrations were studied along a light gradient in the canopies of four temperate deciduous species differing in shade tolerance according to the ranking: Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. = Corylus avellana L. Long-term light environment at the canopy sampling locations was characterized by the fractional penetration of irradiance in the photosynthetically active spectral region (I(sum)). We used a process-based model to distinguish among photosynthesis limitations resulting from variability in fractional nitrogen investments in Rubisco (P(R)), bioenergetics (P(B), N in rate-limiting proteins of photosynthetic electron transport) and light harvesting machinery (P(L), N in chlorophyll and thylakoid chlorophyll-protein complexes). On an area basis, V(cmax) and J(max) (V(a) (cmax) and J(a) (max)) increased with increasing growth irradiance in all species, and the span of variation within species ranged from two (T. cordata) to ten times (C. avellana). Examination of mass-based V(cmax) and J(max) (V(m) (cmax) and J(m) (max)) demonstrated that the positive relationships between area-based quantities and relative irradiance mostly resulted from the scaling of leaf dry mass per area (M(A)) with irradiance. Although V(m) (cmax) and J(m) (max) were positively related to growth irradiance in C. avellana, and J(m) (max) was positively related to irradiance in P. tremula, the variation range was only a factor of two. Moreover, V(m) (cmax) and J(m) (max) were negatively correlated with relative irradiance in T. cordata. Rubisco activity in crude leaf extracts generally paralleled the gas-exchange data, but it was independent of light in T. cordata, suggesting that declining V(m) (cmax) with increasing relative irradiance was related to increasing diffusive resistances from the intercellular air spaces to the sites of carboxylation in this species. Because irradiance had little effect on foliar nitrogen concentration, the relationships of P(B) and P(R) with irradiance were similar to those of V(m) (cmax) and J(m) (max). Shade-intolerant species tended to have greater P(B) and P(R) and also larger V(a) (cmax) and J(a) (max) than more shade-tolerant species. However, for the whole material, P(B) and P(R) varied only about 50%, whereas V(a) (cmax) and J(a) (max) varied more than 15-fold, further emphasizing the importance of leaf anatomical plasticity in determining photosynthetic acclimation to high irradiance. Leaf chlorophyll concentrations and fractional nitrogen investments in light harvesting increased hyperbolically with decreasing irradiance to improve quantum use efficiency for incident irradiance. The effect of irradiance on P(L) was of the same order as its effect in the opposite direction on M(A), leading to either a constant model estimate of leaf absorptance with I(sum) or a slightly positive correlation. We conclude that leaf morphological plasticity is a more relevant determinant of foliage adaptation to high irradiance than foliage biochemical properties, whereas biochemical adaptation to low irradiance is of the same magnitude as the anatomical adjustments. Although shade-tolerant species did not have greater chlorophyll concentrations and P(L) than shade-intolerant species, they possessed lower M(A), and could maintain a more extensive foliar display for light capture with constant biomass investment in leaves.