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PubMed1996

Nutrient retranslocation in the foliage of Pinus sylvestris L. growing along a heavy metal pollution gradient.

Retranslocation of N, P, K, Ca, Mg, Mn, Fe, Zn, Cu and Ni within the foliage of Scots pine (Pinus sylvestris L.) was studied during autumnal needle senescence along a heavy-metal pollution gradient in western Finland. The stands were located at distances of 0.5 (Har 0.5), 4 (Har 4) and 8 km (Har 8) to the southeast of a copper-nickel smelter at Harjavalta. A background study stand was located at Hämeenkangas (Häm), 60 km northeast of the smelter. During needle senescence, the mobile nutrients N, P and K decreased by 67-88% and needle dry mass decreased by 31-43%. Retranslocation of P and K was less efficient in the stand nearest the smelter (Har 0.5) than in the other stands. During needle senescence at stand Har 0.5, heavy metal content either decreased slightly (Zn, Ni) or increased (Fe, Cu), whereas in the background study stand, retranslocation efficiency of Cu was 75%. Net throughfall (the amount of a nutrient in throughfall minus the amount in open precipitation) was higher in stand Har 0.5 than in stand Har 8 for all of the elements measured, except N and P which were intercepted by the canopy. In stand Har 8, deposition of S, Cu and Ni was intercepted by the canopy, and net throughfall of Fe and Zn was very low. In contrast, S and heavy metal contents of net throughfall in stand Har 0.5 were greater than the decrease in these elements in the needles indicating that accumulation on needle surfaces and subsequent wash-off accounted for a major part of the fluxes. Furthermore, the Cu content in net throughfall during the senescence period was higher than the original Cu content of the needles in July, providing additional evidence that much of the needle Cu content was comprised of dry deposits of Cu on the needle surface.

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PubMed1996

Light environment alters ozone uptake per net photosynthetic rate in black cherry trees.

Foliar ozone uptake rates of different-sized black cherry (Prunus serotina Ehrh.) trees were compared within a deciduous forest and adjacent openings in north-central Pennsylvania during one growing season. Study trees included open-grown seedlings and saplings, forest understory seedlings and saplings, and sunlit and shaded portions of mature canopy tree crowns. Instantaneous ozone uptake rates were highest in high-light environments primarily because of higher stomatal conductances. Low ozone uptake rates of seedlings and saplings in the forest understory could be attributed partially to lower average ambient ozone concentrations compared to the canopy and open environments. Among the tree size and light combinations tested, ozone uptake rates were highest in open-grown seedlings and lowest in forest-grown seedlings. Despite lower ozone uptake rates of foliage in shaded environments, ozone uptake per net photosynthesis of foliage in shaded environments was significantly higher than that of foliage in sunlit environments because of weaker coupling between net photosynthesis and stomatal conductance in shaded environments. The potential for greater ozone injury in shaded environments as a result of greater ozone uptake per net photosynthesis is consistent with previous reports of greater ozone injury in shaded foliage than in sunlit foliage.

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PubMedDate not supplied

Transpiration of a boreal pine forest measured by branch bag, sap flow and micrometeorological methods.

Three independent methods were used to evaluate transpiration of a boreal forest: the branch bag, sap flow and eddy covariance methods. The branch bag method encloses several thousand needles and gives a continuous record of branch transpiration. The sap flow method provides a continuous record of sap velocity and an estimate of tree transpiration. The eddy covariance method typically measures evaporation rates between a forest and the atmosphere. We deployed an extra eddy covariance system below the forest to estimate canopy transpiration by difference. The three systems detected small water vapor fluxes despite a plentiful supply of energy to drive evaporation. We also observed that transpiration rates were low even when the soil was well supplied with water. Low rates of transpiration were attributed to the canopy's low leaf area index and the marked reduction in stomatal conductance as vapor pressure deficits increased. Water vapor fluxes, derived from the sap flow method, lagged behind those derived by the branch bag method by 1 to 2 h. The sap flow method also suffered from sampling errors caused by the non-uniformity of flow across the sapwood and the spatial variability of sapwood cross section throughout the forest. Despite technical difficulties associated with hourly measurements, daily totals of transpiration agreed well with values derived from micrometeorological systems.

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PubMedDate not supplied

Regulation of branch-level gas exchange of boreal trees: roles of shoot water potential and vapor pressure difference.

Effects of shoot water potential (Psi) and leaf-to-atmosphere vapor pressure difference (VPD) on gas exchange of jack pine (Pinus banksiana Lamb.), black spruce (Picea mariana (Mill.) B.S.P.), and aspen (Populus tremuloides Michx.) were investigated at the northern edge of the boreal forest in Manitoba, Canada. Laboratory measurements on cut branches showed that net photosynthesis (A(n)) and mesophyll conductance (g(m)) of jack pine and g(m) of black spruce did not respond to Psi until a threshold Psi was reached below which they decreased linearly. Photosynthesis of black spruce decreased slowly with decreasing Psi above the threshold and declined more rapidly thereafter. The threshold Psi was lower in black spruce than in jack pine. However, stomatal conductance (g(s)) of black spruce decreased continuously with decreasing Psi, whereas g(s) of jack pine showed a threshold response. Mesophyll limitations were primarily responsible for the decline in A(n) at low Psi for jack pine and black spruce in the middle of the growing season, but stomatal limitations became more important later in the season. Field measurements on in situ branches on warm sunny days showed that both conifer species maintained Psi above the corresponding threshold and there was no evidence of Psi limitation on A(n) of jack pine, black spruce or aspen. Vapor pressure difference was important in regulating gas exchange in all three species. An empirical model was used to quantify the g(s) response to VPD. When parameterized with laboratory data for the conifers, the model also fit the corresponding field data. When parameterized with field data, the model showed that stomata of aspen were the most sensitive of the three species to VPD, and stomata of black spruce were the least sensitive. For jack pine and aspen, stomata of foliage in the upper canopy were significantly more sensitive than stomata of foliage in the lower canopy. Vapor pressure difference had a greater impact on A(n) of aspen than on A(n) of the conifers as a result of aspen's greater stomatal sensitivity to VPD and greater slope of the relationship between A(n) and intercellular CO(2) concentration (C(i)). During the 1994 growing season, VPD averaged 1.0 kPa, corresponding to ratios of C(i) to ambient CO(2) of 0.77, 0.71 and 0.81 for jack pine, black spruce and aspen, respectively. We conclude that increases in VPD at the leaf surface in response to climate change should affect the absolute CO(2) and H(2)O fluxes per unit leaf area of the aspen component of a boreal forest landscape more than those of the conifer component.

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PubMed1998

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.

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PubMed1998

Net ecosystem productivity, net primary productivity and ecosystem carbon sequestration in a Pinus radiata plantation subject to soil water deficit.

Tree carbon (C) uptake (net primary productivity excluding fine root turnover, NPP') in a New Zealand Pinus radiata D. Don plantation (42 degrees 52' S, 172 degrees 45' E) growing in a region subject to summer soil water deficit was investigated jointly with canopy assimilation (A(c)) and ecosystem-atmosphere C exchange rate (net ecosystem productivity, NEP). Net primary productivity was derived from biweekly stem diameter growth measurements using allometric relations, established after selective tree harvesting, and a litterfall model. Estimates of A(c) and NEP were used to drive a biochemically based and environmentally constrained model validated by seasonal eddy covariance measurements. Over three years with variable rainfall, NPP' varied between 8.8 and 10.6 Mg C ha(-1) year(-1), whereas A(c) and NEP were 16.9 to 18.4 Mg C ha(-1) year(-1) and 5.0-7.2 Mg C ha(-1) year(-1), respectively. At the end of the growing season, C was mostly allocated to wood, with nearly half (47%) to stems and 27% to coarse roots. On an annual basis, the ratio of NEP to stand stem volume growth rate was 0.24 +/- 0.02 Mg C m(-3). The conservative nature of this ratio suggests that annual NEP can be estimated from forest yield tables. On a biweekly basis, NPP' repeatedly lagged A(c), suggesting the occurrence of intermediate C storage. Seasonal NPP'/A(c) thus varied between nearly zero and one. On an annual basis, however, NPP'/A(c) was 0.54 +/- 0.03, indicating a conservative allocation of C to autotrophic respiration. In the water-limited environment, variation in C sequestration rate was largely accounted for by a parameter integrative for changes in soil water content. The combination of mensurational data with canopy and ecosystem C fluxes yielded an estimate of heterotrophic respiration (NPP' - NEP) approximately 30% of NPP' and approximately 50% of NEP. The estimation of fine-root turnover rate is discussed.

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PubMed1998

Storage of foliar-absorbed nitrogen and remobilization for spring growth in young nectarine (Prunus persica var. nectarina) trees.

The effectiveness of spraying foliage with urea to provide nitrogen (N) to augment the seasonal internal cycling of N in young nectarine trees (Prunus persica (L.) Batsch var. nectarina (Ait. f. Maxim.), cv. Stark Red Gold) was studied. One-year-old trees were grown with contrasting N supplies during the summer and foliage was sprayed with a 2% urea solution labeled with (15)N just before leaf senescence started. After leaf abscission had finished, the trees were repotted in sand and given no further N. Remobilization of both labeled and unlabeled N for leaf growth the following spring was quantified. Leaves absorbed between 58 and 69% of the (15)N intercepted by the canopy irrespective of tree N status. During leaf senescence, the majority of (15)N was withdrawn from the leaves into the shoot and roots. Remobilization of (15)N the following spring was also unaffected by tree N status. About 38-46% of (15)N in the trees was recovered in the new growth. More unlabeled N (derived from root uptake) was remobilized for leaf growth in the spring than was withdrawn from leaves during canopy senescence the previous autumn. Therefore, soil-applied N augmented N storage pools directly, and contributed more to N remobilization the following spring than did foliar-absorbed (15)N.

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PubMed1998

The response of Pinus sylvestris to drought: stomatal control of transpiration and hydraulic conductance.

We investigated the impact of drought on the physiology of 41-year-old Scots pine (Pinus sylvestris L.) in central Scotland. Measurements were made of the seasonal course of transpiration, canopy stomatal conductance, needle water potential, xylem water content, soil-to-needle hydraulic resistance, and growth. Comparison was made between drought-treated plots and those receiving average precipitation. In response to drought, transpiration rate declined once volumetric water content (VWC) over the top 20 cm of soil reached a threshold value of 12%. Thereafter, transpiration was a near linear function of soil water content. As the soil water deficit developed, the hydraulic resistance between soil and needles increased by a factor of three as predawn needle water potential declined from -0.54 to -0.71 MPa. A small but significant increase in xylem embolism was detected in 1-year-old shoots. Stomatal control of transpiration prevented needle water potential from declining below -1.5 MPa. Basal area, and shoot and needle growth were significantly reduced in the drought treatment. In the year following the drought, canopy stomatal conductance and soil-to-needle hydraulic resistance recovered. Current-year needle extension recovered, but a significant reduction in basal area increment was evident one year after the drought. The results suggest that, in response to soil water deficit, mature Scots pine closes its stomata sufficiently to prevent the development of substantial xylem embolism. Reduced growth in the year after a severe soil water deficit is most likely to be the result of reduced assimilation in the year of the drought, rather than to any residual embolism carried over from one year to the next.

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PubMedDate not supplied

Passive nighttime warming facility for forest ecosystem research.

A nighttime warming experiment is proposed. Over the last four decades a significant rise in nighttime minimum temperature has been determined from analysis of meteorological records from a global distribution of locations. The experiment involves nighttime deployment of infrared (IR) reflecting curtains around four sides of a forest canopy and across the top of the forest to mimic the top-down warming effect of cloud cover. The curtains are deployed with cable and pulley systems mounted on a tower and scaffolding structure built around the selected forest site. The trunk space is not enclosed except as an optional manipulation. The curtains reflect long-wave radiation emitted from the forest and ground back into the forest warming the trees, litter, and soil. Excellent infrared reflection can be obtained with commercially available fabrics that have aluminum foil bonded to one side. A canopy warming of 3 to 5 degrees C is expected on cloudless nights, and on cloudy nights, a warming of 1 to 3 degrees C is anticipated relative to a control plot. The curtains are withdrawn by computer control during the day and also at night during periods with precipitation or excessive wind. Examples of hypothesized ecosystem responses to nighttime warming include: (1) increase in tree maintenance respiration (decreasing carbon reserves and ultimately tree growth), (2) increase in the length of the growing season (increasing growth), (3) increase in soil respiration, (4) increase in litter decomposition, (5) increase in mineralization of N and other nutrients from soil organic matter, (6) increase in nutrient uptake (increasing growth), and (7) increase in N immobilization in litter. Hypothesis 1 has the opposite consequence for tree growth to Hypotheses 2 and 6, and thus opposite consequences for the feedback regulation that vegetation has on net greenhouse gas releases to the atmosphere. If Hypothesis 1 is dominant, warming could lead to more warming from the additional CO(2) emissions. Site-specific meteorological, ecophysiological, and phenological measurements are obtained in the warming treatment and in a carefully selected control plot to investigate site-specific hypotheses. Measurements made on both plots for a baseline period and during the period of curtain deployment provide data to test the hypotheses statistically by the "before-after-control-impact" method applicable to unreplicated experiments. The enclosure has a modular design that can be adapted and combined with other forest-scale manipulation experiments such as free air CO(2) enrichment and throughfall displacement.

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PubMed1999

Modeling the effect of physiological responses to green pruning on net biomass production of Eucalyptus nitens.

Green pruning of Eucalyptus nitens (Deane and Maiden) Maiden increases instantaneous rates of light-saturated CO(2) assimilation (A), and changes patterns of total leaf area and foliage distribution. We investigated the importance of such changes on the rate of recovery of growth following pruning. A simple process-based model was developed to estimate daily net biomass production (G(d)) of three-year-old plantation-grown trees over a 20-month period. The trees had been pruned by removal of 0, 50 or 70% of the length of green crown, equivalent to removal of 0, 55 or 88% of leaf area, respectively, when the plantation verged on canopy closure. Total G(d) was reduced by only 20% immediately following the 50%-pruning treatment, as a result of both the high leaf dark respiration and low A in the portion of the crown removed compared to the top of the crown. Pruning at the time of canopy closure preempted a natural and rapid decline in G(d) of the lower crown. Although leaf area index (L) was approximately 6.0 at the time of pruning, high light interception (95%) occurred with an L of 4.0. The 50%-pruning treatment reduced L to 3.5, but the physiological responses to pruning were sufficient to compensate fully for the reduction in intercepted radiation within 110 days of pruning. The 70%-pruning treatment reduced L to 1.9, and reduced G(d) by 77%, reflecting the removal of branches with high A in the mid and upper crown. Physiological responses to the 70%-pruning treatment were insufficient to increase G(d) to the value of unpruned trees during the study. Model sensitivity analysis showed that increases in A following pruning increased G(d) by 20 and 25% in the 50- and 70%-pruned trees, respectively, 20 months after pruning. Changes in leaf area/foliage distribution had a greater effect on G(d) of 50%-pruned trees (47% increase) than did changes in A. However, the reduction in photosynthetic potential associated with the 70%-pruning treatment resulted in only small changes in leaf area/foliage distribution, which consequently had little effect on G(d). The effects of physiological processes occurring within the crown and in response to green pruning on G(d) are discussed with respect to pruning of plantations.

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PubMed1999

Differences in chemical composition relative to functional differentiation between petioles and laminas of Fraxinus excelsior.

Differences in structural and nonstructural carbohydrates, lignin and chlorophyll, and Rubisco (ribulose-1,5-bisphosphate carboxylase/oxygenase) activity between petioles and leaflets were studied along a canopy light gradient in Fraxinus excelsior L., which has pinnate compound leaves and up to 20% of foliar biomass invested in petioles. Long-term light conditions at the sampling locations were characterized by values of seasonal mean integrated quantum flux density (Q(int), mol m(-2) day(-1)) estimated by combining data from hemispherical photographs at the sampling locations with measurements of global solar radiation above the canopy during the growing season. The contribution of petioles to leaf carbon assimilation was disproportionally lower than that of leaf laminas. Though the light relationships of assimilative compounds-foliar chlorophyll concentration increasing with decreasing Q(int) to improve leaf absorptance, foliar N concentration and Rubisco activity being relatively constant along the light gradient-were similar for both petioles and leaflets, petiole nitrogen and chlorophyll concentrations were only 30% and 10%, respectively, of those of leaflets. Nonstructural carbohydrate concentration was about 20% higher in petioles than in leaf laminas, indicating that petioles also serve as storage tissues for photosynthates. Relationships between foliar structural carbon components and irradiance-increasing lignin (L) and decreasing structural polysaccharide (SP) concentrations with increasing Q(int)-were qualitatively similar for petioles and leaflets. However, petioles had lower L, but higher SP and total investment in structural compounds (L + SP) than leaflets. Greater lignification at high irradiances in leaflets than in petioles was attributed to greater water stresses at high light, and to more variable water contents of actively transpiring leaflets. Low lignin concentration in combination with high osmotically active carbohydrate concentrations in petioles suggest that turgor plays an important role in the mechanical properties of petioles. As a result of lower lignin and protein concentrations, the glucose cost of petiole construction (g glucose per g dry mass) was about 5% lower than that of leaf laminas.

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PubMed1999

Seasonal variations in leaf area index, leaf chlorophyll, and water content; scaling-up to estimate fAPAR and carbon balance in a multilayer, multispecies temperate forest.

Seasonal differences in phenology between coniferous and deciduous tree species need to be considered when developing models to estimate CO(2) exchange in temperate forest ecosystems. Because seasonal variations in CO(2) flux in temperate forests are closely correlated with plant phenology, we quantified the phenology of forest species in a multilayered forest with patches of Scots pine (Pinus sylvestris L.) and oak (Quercus robur L.) in Brasschaat, Belgium. A scaling-up modeling approach was developed to simulate reflectance at the leaf and canopy scales over a one-year cycle. Chlorophyll concentration, water content, specific leaf area and leaf area index of the forest species were measured throughout an entire year (1997). Scaling-up from the leaf to canopy was achieved by linking the PROSPECT and SAIL models. The result is the annual progression of the fraction of absorbed photosynthetically active radiation (fAPAR) in a 1 km(2) forest area, which can be directly related to high-resolution, remotely sensed data.

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PubMed1999

A micro-power precision amplifier for converting the output of light sensors to a voltage readable by miniature data loggers.

To record photosynthetically active radiation (PAR) simultaneously at a number of points throughout a forest canopy, we developed a simple, inexpensive (< $10 US) current-to-voltage converter that processes the current generated by a photodiode radiation sensor to a voltage range that is recordable with a miniature data logger. The converter, which weighs less than 75 g and has a volume of only 100 cm(3), is built around an ultra-low power OP-90 precision operational amplifier, which consumes less than 0.5 mA at 9 V when converting the output of a Li-Cor LI-190SA quantum sensor exposed to photosynthetically active radiation (PAR) of 2500 &mgr;mol m(-2) s(-1) or only 5 &mgr;A in low light. A small 9-V battery thus powers the amplifier for more than 1000 h of continuous operation. Correlations between photometer readings and voltage output from the current-to-voltage converter were high and linear at both high and low PAR. Sixteen Li-Cor LI-190SA quantum sensors each equipped with current-to-voltage converters and connected to a miniature data logger were deployed in the upper branches of a Panamanian tropical rainforest canopy. Each unit performed reliably during a one- or two-week evaluation.

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PubMed1986

Coupling of carbon and water interactions in forest stands.

Some measured values of the ratio transpiration rate/assimilation rate (E/A) for a Sitka spruce stand in Scotland and an oak-hickory stand are presented and compared with E/A for some agricultural crops. Typical midday values of E/A (on a molar basis) range from 100 to 900. Analysis suggests that this variation can be explained by differences in climate, canopy conductance and degree of coupling of the vegetation to the atmosphere. During the day, E/A varied considerably in the tree stands at the same time as there were changes in radiation, vapor saturation deficit and temperature. A lumped parameter, 'big leaf' model of canopy transpiration and assimilation was used to provide insight into how this complex of variables might influence E/A. The model predicted little response to radiation, but increase in E/A with increasing vapor saturation deficit and temperature, in agreement with the trends in the data. It was concluded that there is a great need for measurements of E/A of stands in relation to nutrient and water stress.

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PubMed1986

Carbon x nutrient interactions-the limitations to productivity.

The interaction of nutrient cycles with the production, accumulation and death of organic matter is considered, in particular the relation to changes with stand age. It is shown that, because of efficient recycling within the tree and through the litter, the demands made by forests on the soil nutrient capital fall markedly after canopy closure. Because this effect is primarily a result of the ability of the tree to conserve and reuse nutrients in the foliage, the significance of the evergreen habit is examined. Although there are many reports of a net photosynthetic contribution from foliage older than one year, the relationships examined suggest that whereas there is virtually no correlation between growth rate and total foliage carried, there is quite a good relationship with weight of current-year foliage irrespective of whether trees are deciduous or the age at which evergreen foliage is shed. It is suggested that the evergreen habit would be a nutritional disadvantage prior to canopy closure but thereafter a considerable advantage.

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PubMed2000

Forest growth and species distribution in a changing climate.

Climate change has many potential effects on plants, some detrimental to growth, others beneficial. Increasing CO(2) concentration can increase photosynthetic rates, with the greatest increases likely to be in C(3) plants growing in warm dry conditions. Increasing temperature directly affects plant growth through effects on photosynthetic and respiration rates. However, plants have a considerable ability to adapt to changing conditions and can tolerate extremely high temperatures, provided that adequate water is available. Increasing temperature may increase vapor pressure deficits of the air, and thereby increase transpiration rates from most plant canopies. Effects are likely to vary among plant communities, with forests generally experiencing greater increases in transpiration rates than grasslands. These increases in transpiration are likely to be reduced by stomatal closure in response to increasing CO(2) concentration. In many areas, precipitation will probably increase with global warming; however, these increases may be insufficient to meet the increased transpirational demand by plant canopies. Increasing temperature is likely to increase soil organic matter decomposition rates so that nutrients may be more readily mineralized and made available to plants. In highly fertile systems, this could lead to nutrient losses through leaching. For different combinations of increases in temperature and CO(2) concentration, and for systems primarily affected by water or nutrient limitations, different overall effects on plant productivity can be expected. Responses will be negative in some circumstances and positive in others, but on the whole, catastrophic changes to forest growth seem unlikely under most conditions. In contrast,ecological consequences of climate change are potentially more serious. The distribution of many species tends to be limited to a narrow range of environmental conditions. Climate conditions over much of a species' current natural range may therefore become unsuitable, leading to significant decline of forests or of particular species within forests.

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PubMed2000

Spatial and seasonal variability of photosynthetic parameters and their relationship to leaf nitrogen in a deciduous forest.

We used gas exchange techniques to estimate maximum rate of carboxylation (V(cmax)), a measure of photosynthetic capacity, in the understory and upper crown of a closed deciduous forest over two seasons. There was extensive variability in photosynthetic capacity as a result of vertical canopy position, species type, leaf age and drought. Photosynthetic capacity was greater in oaks than in maples and greater in the overstory than in the understory. Parameter V(cmax) was maximal early in the season but declined slowly throughout most of the summer, and then more rapidly during senescence. There was also an apparent decline during drought in some trees. Variability in V(cmax) as a result of species or vertical canopy gradients was described well by changes in leaf nitrogen per unit area (N(a)). However, temporal changes in V(cmax) were often poorly correlated with leaf nitrogen, especially in spring and summer and during drought. This poor correlation may be the result of a seasonally dependent fractional allocation of leaf nitrogen to Rubisco; however, we could not discount Rubisco inactivation, patchy stomatal closure or changes in mesophyll resistance. Consequently, when a single annual regression equation of V(cmax) versus N(a) was used for this site, there were substantial errors in the temporal patterns in V(cmax) that will inevitably result in modeling errors.

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PubMed2001

A theoretical analysis of the influence of heterogeneity in chlorophyll distribution on leaf reflectance.

Attempts to determine the vitality of vegetation and to detect vegetation stress from remotely sensed data have focused on chlorophyll concentration, because it influences the reflectance of vegetation and tends to correlate with vegetation health and stress. Pollution, pathogens and pests can cause localized regions of chlorosis and necrosis across a leaf surface, but the extent to which these patches influence the overall reflectance and spectral signature of the leaf and canopy has not been tested. A conifer leaf model (LIBERTY), which simulates the influence of leaf biochemical concentrations of chlorophyll, water, lignin, cellulose and protein on the reflectance of leaves from 400 to 2500 nm, was used to determine the effect of patches of chlorosis on leaf reflectance. A fraction of the leaf f is assumed to be chlorotic with a chlorophyll concentration C(1). The remainder of the leaf has chlorophyll concentration C(2) such that mean leaf chlorophyll concentration, C(mean) = fC(1) + (1 - f)C(2), is constant for a range of f and C(1) values. LIBERTY can be used to estimate the reflectance of a leaf with a particular chlorophyll concentration at a particular wavelength R(lambda,C) (assuming other leaf properties remain constant), thus we can estimate the reflectance of the chlorotic leaf as fR(lambda,C(1))+ (1 - f)R(lambda,C(2)). The model indicated that small areas of chlorosis have a disproportionately large influence on overall leaf reflectance. For example, a leaf with 25% of its area chlorotic can have the same reflectance (400-700 nm) as a homogeneous leaf with 60% less chlorophyll. Thus, determination of chlorophyll concentration from remotely sensed data is prone to underestimation when chlorophyll is nonuniformly distributed. Hence, attempts to model leaf and canopy reflectance using radiative transfer models will need to consider how to incorporate nonuniform chlorophyll distribution.

Chlorophyll↗