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Effects of light availability and tree size on the architecture of assimilative surface in the canopy of Picea abies: variation in needle morphology.

Needle dimensions, needle surface area, needle dry weight per area (LWA) and needle density (ND, needle weight per volume) were measured in terminal current-year shoots in a natural canopy of variably sized Picea abies (L.) Karst. trees growing along a light gradient. Needle shape was described as a rhomboid. Needle width (D(2)) increased with increasing diffuse site factor, a(d) (relative amount of penetrating diffuse solar radiation), whereas needle thickness (D(1)) remained nearly constant, resulting in an inverse relationship between D(1)/D(2) and a(d) and an increase in the ratio of total (TLA) to projected needle surface area (PLA) with increasing a(d). Because of the variations in needle morphology with respect to light availability, the shoot parameters used in present canopy models are also expected to be light-sensitive, and studies involving shoot morphology should also consider the variability in needle geometry. Needle dimensions and total tree height were not correlated. However, LWA increase with both increasing a(d) and total tree height. When LWA was expressed as the product of ND and needle height (NH, height of the rhomboidal transverse section of a needle), LWA appeared to increase with irradiance, because of changing NH, and with total tree height, because of changing needle density.

Journal Article↗

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

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

Journal Article↗

Canopy conductance of Pinus taeda, Liquidambar styraciflua and Quercus phellos under varying atmospheric and soil water conditions.

Sap flow, and atmospheric and soil water data were collected in closed-top chambers under conditions of high soil water potential for saplings of Liquidambar styraciflua L., Quercus phellos L. and Pinus taeda L., three co-occurring species in the southeastern USA. Responses of canopy stomatal conductance (g(t)) to water stress induced by high atmospheric water vapor demand or transpiration rate were evaluated at two temporal scales. On a diurnal scale, the ratio of canopy stomatal conductance to maximum conductance (g(t)/g(t,max)) was related to vapor pressure deficit (D), and transpiration rate per unit leaf area (E(l)). High D or E(l) caused large reductions in g(t)/g(t,max) in L. styraciflua and P. taeda. The response of g(t)/g(t,max) to E(l) was light dependent in L. styraciflua, with higher g(t)/g(t,max) on sunny days than on cloudy days. In both L. styraciflua and Q. phellos, g(t)/g(t,max) decreased linearly with increasing D (indicative of a feed-forward mechanism of stomatal control), whereas g(t)/g(t,max) of P. taeda declined linearly with increasing E(l) (indicative of a feedback mechanism of stomatal control). Longer-term responses to depletion of soil water were observed as reductions in mean midday g(t)/g(t,max), but the reductions did not differ significantly between species. Thus, species that employ contrasting methods of stomatal control may show similar responses to soil water depletion in the long term.

Journal Article↗

Regulation of stomatal conductance and transpiration in forest canopies.

Processes regulating stomatal conductance, g(s), and transpiration, E, from forest canopies are reviewed. The first section deals with the response of g(s) to environmental variables. Phenomenological models have been used to interpret field data and predict diurnal and seasonal variability in g(s), but models that couple stomatal conductance to photosynthesis at the leaf scale are now being used more widely. The vertical distribution of foliar nitrogen concentration is helpful for scaling these processes from leaves to canopies, and the analysis of data from many studies has led to the emergence of simplified, general relationships for estimating evaporation and carbon uptake by forests at stand and regional scales. Evidence for the regulation of stomatal conductance by hydraulic and chemical signals is presented in the second section. Rapid and reversible changes in g(s) following a perturbation to the water potential gradient in the flow pathway suggest that stomata respond directly to hydrostatic signals. Other evidence supports the contention that signals are transmitted by abscisic acid (ABA), possibly originating in the roots. For large woody plants, the short-term responses of stomata are probably brought about by hydraulic signals that affect g(s) by triggering the release of ABA in the leaves. Tardieu and Davies (1993) developed an interactive model that incorporates hydraulic and chemical effects to describe the response of stomata to soil drying and evaporative demand. In the third section, evidence is presented that short-term changes in g(s) are linked closely to the hydraulic properties of the conducting system to minimize loss of hydraulic conductivity through xylem by cavitation. Examples of homeostatic mechanisms that operate to ensure the long-term balance between evaporative demand and the potential hydraulic conductivity of trees growing in different environments are described. Two hypotheses are examined: (1) height growth in trees is limited by the capacity of the conducting system; and (2) the decline in productivity with stand age is attributable to a decrease in conductivity.

Journal Article↗

Effects of CO2 enrichment on the photosynthetic light response of sun and shade leaves of canopy sweetgum (Liquidambar styraciflua) in a forest ecosystem.

To investigate whether sun and shade leaves respond differently to CO2 enrichment, we examined photosynthetic light response of sun and shade leaves in canopy sweetgum (Liquidambar styraciflua L.) trees growing at ambient and elevated (ambient + 200 microliters per liter) atmospheric CO2 in the Brookhaven National Laboratory/Duke University Free Air CO2 Enrichment (FACE) experiment. The sweetgum trees were naturally established in a 15-year-old forest dominated by loblolly pine (Pinus taeda L.). Measurements were made in early June and late August 1997 during the first full year of CO2 fumigation in the Duke Forest FACE experiment. Sun leaves had a 68% greater leaf mass per unit area, 63% more leaf N per unit leaf area, 27% more chlorophyll per unit leaf area and 77% greater light-saturated photosynthetic rates than shade leaves. Elevated CO2 strongly stimulated light-saturated photosynthetic rates of sun and shade leaves in June and August; however, the relative photosynthetic enhancement by elevated CO2 for sun leaves was more than double the relative enhancement of shade leaves. Elevated CO2 stimulated apparent quantum yield by 30%, but there was no interaction between CO2 and leaf position. Daytime leaf-level carbon gain extrapolated from photosynthetic light response curves indicated that sun leaves were enhanced 98% by elevated CO2, whereas shade leaves were enhanced 41%. Elevated CO2 did not significantly affect leaf N per unit area in sun or shade leaves during either measurement period. Thus, the greater CO2 enhancement of light-saturated photosynthesis in sun leaves than in shade leaves was probably a result of a greater amount of nitrogen per unit leaf area in sun leaves. A full understanding of the effects of increasing atmospheric CO2 concentrations on forest ecosystems must take account of the complex nature of the light environment through the canopy and how light interacts with CO2 to affect photosynthesis.

Journal Article↗

Responses of foliar photosynthetic electron transport, pigment stoichiometry, and stomatal conductance to interacting environmental factors in a mixed species forest canopy.

We studied limitations caused by variations in leaf temperature and soil water availability on photosynthetic electron transport rates calculated from foliar chlorophyll fluorescence analysis (U) in a natural deciduous forest canopy composed of shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. In both species, there was a positive linear relationship between light-saturated U (Umax) per unit leaf area and mean seasonal integrated daily quantum flux density (Ss, mol per square m per day). Acclimation of leaf dry mass per area and nitrogen per area to growth irradiance largely accounted for this positive scaling. However, the slopes of the Umax versus Ss relationships were greater on days when leaf temperature was high than on days when leaf temperature was low. Overall, Umax varied 2.5-fold across a temperature range of 20-30 degrees C. Maximum stomatal conductance (Gmax) also scaled positively with Ss. Although Gmax observed during daily time courses, and stomatal conductances during Umax measurements declined in response to seasonally decreasing soil water contents, was insensitive to prolonged water stress, and was not strongly correlated with stomatal conductances during its estimation. These results suggest that photorespiration was an important electron sink when intercellular CO2 concentration was low because of closed stomata. Given that xanthophyll cycle pool size (VAZ, sum of violaxanthin, antheraxanthin, and zeaxanthin) may play an important role in dissipation of excess excitation energy, the response of VAZ to fluctuating light and temperature provided another possible explanation for the stable Umax. Xanthophyll cycle carotenoids per total leaf chlorophyll (VAZ/Chl) scaled positively with integrated light and negatively with daily minimum air temperature, whereas the correlation between VAZ/Chl and irradiance was best with integrated light averaged over 3 days preceding foliar sampling. We conclude that the potential capacity for electron transport is determined by long-term acclimation of U to certain canopy light conditions, and that the rapid adjustment of the capacity for excitation energy dissipation plays a significant part in the stabilization of this potential capacity. Sustained high capacity of photosynthetic electron transport during stress periods provides an explanation for the instantaneous response of U to short-term weather fluctuations, but also indicates that U restricts potential carbon gain under conditions of water limitation less than does stomatal conductance.

Journal Article↗

Seasonal CO(2) assimilation and stomatal limitations in a Pinus taeda canopy.

Net CO(2) assimilation (A(net)) of canopy leaves is the principal process governing carbon storage from the atmosphere in forests, but it has rarely been measured over multiple seasons and multiple years. I measured midday A(net) in the upper canopy of maturing loblolly pine (Pinus taeda L.) trees in the piedmont region of the southeastern USA on 146 sunny days over 36 months. Concurrent data for leaf conductance and photosynthetic CO(2) response curves (A(net)-C(i) curves) were used to estimate the relative importance of stomatal limitations to CO(2) assimilation in the field. In fully expanded current-year and 1-year-old needles, midday light-saturated A(net) was constant over much of the growing season (5-6 &mgr;mol CO(2) m(-2) s(-1)), except during drought periods. During the winter season (November-March), midday A(net) of overwintering needles varied in proportion to leaf temperature. Net CO(2) assimilation at light saturation occurred when daytime air temperatures exceeded 5-6 degrees C, as happened on more than 90% of the sunny winter days. In both age classes of foliage, winter carbon assimilation accounted for approximately 15% of the daily carbon assimilation on sunny days throughout the year, and was relatively insensitive to year-to-year differences in temperature during this season. However, strong stomatal limitations to A(net) occurred as a result of water stress associated with freezing cycles in winter. During the growing season, drought-induced water stress produced the largest year-to-year differences in seasonal CO(2) assimilation on sunny days. Seasonal A(net) was more drought sensitive in current-year needles than in 1-year-old needles. Relative stomatal limitations to daily integrated A(net) were approximately 40% over the growing season, and summer drought rather than high temperatures had the largest impact on summer A(net) and integrated annual CO(2) uptake in the upper crown. Despite significant stomatal limitations, a long duration of near-peak A(net) in the upper crown, particularly in 1-year-old needles, conferred high seasonal and annual carbon gain.

Journal Article↗

Canopy stomatal conductance and xylem sap abscisic acid (ABA) in mature Scots pine during a gradually imposed drought.

We investigated the effect of drought on canopy stomatal conductance (g(c)), and examined the hypothesis that g(c) is controlled by the chemical messenger abscisic acid (ABA) produced in roots. Beginning in November 1994, we subjected a mature stand of Scots pine (Pinus sylvestris L.) to an imposed 11-month drought. Control plots were maintained at average-season soil water content. Xylem sap was extracted from shoots at regular intervals from April to November 1995. Soil water, sap flow and leaf water potentials (predawn to dusk) were recorded at the same time. Canopy stomatal conductance was calculated from sap flow data and xylem sap ABA concentrations ([ABA(xyl)]) were measured by radioimmunoassay. Mean [ABA(xyl)] in control trees was 250 micromol m(-3). No diurnal variation in [ABA(xyl)] was detected. With soil drying, [ABA(xyl)] increased to a maximum in summer (600 micromol m(-3)), but decreased again toward autumn; however, no significant increase in ABA flux to the leaves occurred. A decline in g(c) was detected when volumetric soil water content declined below 0.12. The decline in g(c) could not have been mediated by increasing [ABA(xyl)] because stomatal closure appeared to precede any increase in [ABA(xyl)]. Peak sap flow velocity data were used to estimate delivery times for root-to-shoot signals in 15-m tall trees. Under normal field conditions, a signal would take 12 days to travel from the site of production (roots) to the presumed site of action (shoots). However, under drought conditions it may take a chemical signal in excess of 6 weeks. We conclude that a feedforward model of short-term stomatal response to soil drying, based solely on the action of a chemical messenger from the roots, is not applicable in mature conifer trees because signal transmission is too slow.

Abscisic Acid↗

Photosynthesis and respiration of black spruce at three organizational scales: shoot, branch and canopy.

To gain insight into the function of photosynthesis and respiration as processes operating within a global ecosystem, we measured gas exchange of mature black spruce (Picea mariana (Mill.) B.S.P.) trees at three organizational scales: individual shoots, whole branches and a forest canopy. A biochemical model was fitted to these data, and physiological parameters were extracted. Pronounced seasonal variation in the estimated model parameters was found at all three organizational scales, highlighting the need to make physiological measurements throughout the year. For example, it took over 100 days for physiological activity to increase from zero during the springtime thaw to its yearly maximum. Good agreement was found between parameter values estimated for the different organizational scales, suggesting that, in the case of aerodynamically rough, largely mono-specific forest canopies, physiological parameters can be estimated from eddy covariance flux measurements. The small differences between photosynthetic parameters estimated at the different scales suggest that the overall spatial organization of photosynthetic capacity is nearly optimized for carbon uptake at each scale.

Carbon Dioxide↗

Chlorophyll content in eucalypt vegetation at the leaf and canopy scales as derived from high resolution spectral data.

The physiological status of forest canopy foliage is influenced by a range of factors that affect leaf pigment content and function. Recently, several indices have been developed from remotely sensed data that attempt to provide robust estimates of leaf chlorophyll content. These indices have been developed from either hand-held spectroradiometer spectra or high spectral resolution (or hyperspectral) imagery. We determined if two previously published indices (Datt 1999), which were specifically developed to predict chlorophyll content in eucalypt vegetation by remote sensing at the leaf scale, can be extrapolated accurately to the canopy. We derived the two indices from hand-held spectroradiometer data of eucalypt leaves exhibiting a range of insect damage symptoms. We also derived the indices from spectra obtained from high spectral and spatial resolution Compact Airborne Spectrographic Imager 2 (CASI-2) imagery to determine if reasonable estimates at a scale of < 1 m can be achieved. One of the indices (R 850/R 710 index, where R is reflectance) derived from hand-held spectroradiometer data showed a moderate correlation with relative leaf chlorophyll content (r = 0.59, P < 0.05) for all dominant eucalypt species in the study area. The R (850)/R (710) index derived from CASI-2 imagery yielded slightly lower correlations over the entire data set (r = 0.42, P < 0.05), but correlations for individual species were high (r = 0.77, P < 0.05). A scaling analysis indicated that the R (850)/R (710) index was strongly affected by soil and water cover types when pixels were mixed, but appeared to be invariant to changes in proportions of understory, which may limit its application.

Chlorophyll↗

Sources of variation in ecophysiological parameters in Douglas-fir and grand fir canopies.

Forest process models predict ecosystem responses from climate variables and physiological parameters. The parameters describe key ecosystem attributes, often as lumped averages. However, the sources and magnitude of variation in these physiological parameters are unknown, which complicates sampling if models are to be parameterized with field measurements. We measured several key parameters, which had been identified by sensitivity analyses of three models, in Abies grandis (Dougl.) Lindl. and Pseudotsuga menziesii var. glauca (Beissn.) Franco trees throughout the growing season. Trees were sampled at eight sites across the interior northwest of the USA. At each site, fertilized and control plots were sampled. The design provided statistical replication for the analysis of variance within a site, allowing us to draw inferences about a regional population of stands. Specific leaf area (SLA) varied by canopy position and treatment (P = 0.0003), by date of sampling (P < 0.0001) and by species (P = 0.0188). Mass-based foliar nitrogen concentration (%N) increased during the summer in both species (P = 0.0019), but at a faster rate in P. menziesii var. glauca than in A. grandis. Sun foliage had a higher mean %N (1.00, SE = 0.02%) than shade foliage (0.92 +/- 0.01%). Apparent quantum yield (Phi) varied among treatments, between species and between canopy positions; each of these variables interacted with date of sampling (P = 0.0207, P < 0.0001 and P = 0.0344, respectively). In A. grandis, mean Phi values (+/- SE) were 0.048 +/- 0.006 mol CO2 (mol incident photon)-1 for sun foliage and 0.067 +/- 0.007 mol CO2 (mol incident photon)(-1) for shade foliage. In P. menziesii, the corresponding mean Phi values were 0.032 +/- 0.003 and 0.047 +/- 0.004 mol CO2 (mol incident photon)-1. Parameters SLA, %N and Phi were all influenced by date, fertilizer treatment, species and crown position. We discuss methods of inferring quantum yields from light response curves and their utility for parameterizing process models. Parameter mean values are presented for each site; these tables provide a documented data set for the parameterization of models describing the western interior forests of the USA.

Cell Respiration↗

Interspecific variation of photosynthesis and leaf characteristics in canopy trees of five species of Dipterocarpaceae in a tropical rain forest.

Photosynthetic rate, nitrogen concentration and morphological properties of canopy leaves were studied in 18 trees, comprising five dipterocarp species, in a tropical rain forest in Sarawak, Malaysia. Photosynthetic rate at light saturation (Pmax) differed significantly across species, varying from 7 to 18 micro mol m(-2) s(-1). Leaf nitrogen concentration and morphological properties, such as leaf blade and palisade layer thickness, leaf mass per area (LMA) and surface area of mesophyll cells per unit leaf area (Ames/A), also varied significantly across species. Among the relationships with leaf characteristics, Pmax had the strongest correlation with leaf mesophyll parameters, such as palisade cell layer thickness (r2 = 0.76, P < 0.001) and Ames/A (r2 = 0.73, P < 0.001). Leaf nitrogen concentration and Pmax per unit area also had a significant but weaker correlation (r2 = 0.46, P < 0.01), whereas Pmax had no correlation, or only weakly significant correlations, with leaf blade thickness and LMA. Shorea beccariana Burck, which had the highest P(max) of the species studied, also had the thickest palisade layer, with up to five or more layers. We conclude that interspecific variation in photosynthetic capacity in tropical rain forest canopies is influenced more by leaf mesophyll structure than by leaf thickness, LMA or leaf nitrogen concentration.

Ericales↗

Leaf longevity of western red cedar (Thuja plicata) increases with depth in the canopy.

Leaf longevity determines the annual cost of replacing foliage biomass and influences water and nutrient budgets. Longevity is readily estimated in most evergreen species by annual bud scars along the shoot. However, some species with indeterminate growth do not show these annual markers, making estimation of longevity difficult. One of these species is the widespread and economically valuable western red cedar (Thuja plicata J. Donn ex D. Don), for which no dependable estimates of leaf longevity exist. In this study, we estimated leaf longevity for western red cedar by counting growth rings in shoots at the point of leaf abscission. Estimates were obtained on 26 dominant or codominant trees growing in natural stands in a montane forest in northern Idaho, USA. Leaf longevity averaged 8.9 (SE = 0.2) years, but it strongly increased with depth in the canopy (0.3 year m(-1); mean crown depth was 15 m), increasing from a mean of 6.8 years in the upper third of the canopy to 10.6 years in the lower third. The increase in longevity with depth in the crown is consistent with many reports showing that longevity increases in resource-limiting environments. Longevity did not vary significantly with altitude or solar insolation in these montane forests. Among stand-level variables, longevity was correlated only with leaf area index: it increased slightly in stands with high leaf area indices. This approach to longevity estimation may be useful for any species that produces annual rings but no obvious bud scars, including many Cupressaceae species.

Altitude↗

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand.

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.

Biomass↗

Influences of canopy photosynthesis and summer rain pulses on root dynamics and soil respiration in a young ponderosa pine forest.

Our first objective was to link the seasonality of fine root dynamics with soil respiration in a ponderosa pine (Pinus ponderosa P. & C. Lawson) plantation located in the Sierra Nevada of California. The second objective was to examine how canopy photosynthesis influences fine root initiation, growth and mortality in this ecosystem. We compared CO2 flux measurements with aboveground and belowground root dynamics. Initiation of fine root growth coincided with tree stem thickening and shoot elongation, preceding new needle growth. In the spring, root, shoot and stem growth occurred simultaneously with the increase in canopy photosynthesis. Compared with the other tree components, initial growth rate of fine roots was the highest and their growing period was the shortest. Both above and belowground components completed 90% of their growth by the end of July and the growing season lasted approximately 80 days. The period for optimal growth is short at the study site because of low soil temperatures during winter and low soil water content during summer. High photosynthetic rates were observed following unusual late-summer rains, but tree growth did not resume. The autotrophic contribution to soil respiration was 49% over the whole season, with daily contributions ranging between 18 and 87%. Increases in soil and ecosystem respiration were observed during spring growth; however, the largest variation in soil respiration occurred during summer rain events when no growth was observed. Both the magnitude and persistence of the soil respiration pulses were positively correlated with the amount of rain. These pulses accounted for 16.5% of soil respiration between Days 130 and 329.

Carbon Dioxide↗

Species-specific bird functions in a forest-canopy food web.

Bird functions in a forest-canopy food web were evaluated by a large-scale field experiment using 'canopy' enclosures. By controlling the presence of two bird species, great tits (Parus major; foliage gleaner) and nuthatches (Sitta europaea; trunk gleaner), in the enclosures, their effect on predatory insects (ants), herbivorous insects (Lepidoptera larvae) and producers (oak trees) was quantified. Great tits reduced the density of Lepidoptera larvae and, indirectly, leaf damage, but had no impact on ants. Nuthatches decreased the density of ants but did not influence either Lepidoptera larvae or leaf damage. These results highlight species-specific functions of birds in the maintenance of forest ecosystems.

Animals↗

Estimation of leaf nitrogen content from spectral characteristics of rice canopy.

Ground-based remotely sensed reflectance spectra of hyperspectral resolution were monitored during the growing period of rice under various nitrogen application rates. It was found that reflectance spectrum of rice canopy changed in both wavelength and reflectance as the plants developed. Fifteen characteristic wavebands were identified from the apparent peaks and valleys of spectral reflectance curves, in accordance with the results of the first-order differentiation, measured over the growing season of rice. The bandwidths and center wavelengths of these characteristic wavebands were different among nitrogen treatments. The simplified features by connecting these 15 characteristic wavelengths may be considered as spectral signatures of rice canopy, but spectral signatures varied with developmental age and nitrogen application rates. Among these characteristic wavebands, the changes of the wavelength in band 11 showed a positive linear relationship with application rates of nitrogen fertilizer, while it was a negative linear relationship in band 5. Mean reflectance of wavelengths in bands 1, 2, 3, 5, 11, and 15 was significantly correlated with application rates. Reflectance of these six wavelengths changed nonlinearly after transplanting and could be used in combination to distinguish rice plants subjected to different nitrogen application rates. From the correlation analyses, there are a variety of correlation coefficients for spectral reflectance to leaf nitrogen content in the range of 350-2400 nm. Reflectance of most wavelengths exhibited an inverse correlation with leaf nitrogen content, with the largest negative value (r = -0.581) located at about 1376 nm. Changes in reflectance at 1376 nm to leaf nitrogen content during the growing period were closely related and were best fitted to a nonlinear function. This relationship may be used to estimate and to monitor nitrogen content of rice leaves during rice growth. Reflectance of red light minimum and near-infrared peak and leaf nitrogen content were correlated nonlinearly.

Agriculture↗

Canopy CO2 enrichment permits tracing the fate of recently assimilated carbon in a mature deciduous forest.

How rapidly newly assimilated carbon (C) is invested into recalcitrant structures of forests, and how closely C pools and fluxes are tied to photosynthesis, is largely unknown. A crane and a purpose-built free-air CO2 enrichment (FACE) system permitted us to label the canopy of a mature deciduous forest with 13C-depleted CO2 for 4 yr and continuously trace the flow of recent C through the forest without disturbance. Potted C4 grasses in the canopy ('isometers') served as a reference for the C-isotope input signal. After four growing seasons, leaves were completely labelled, while newly formed wood (tree rings) still contained 9% old C. Distinct labels were found in fine roots (38%) and sporocarps of mycorrhizal fungi (62%). Soil particles attached to fine roots contained 9% new C, whereas no measurable signal was detected in bulk soil. Soil-air CO2 consisted of 35% new C, indicating that considerable amounts of assimilates were rapidly returned back to the atmosphere. These data illustrate a relatively slow dilution of old mobile C pools in trees, but a pronounced allocation of very recent assimilates to C pools of short residence times.

Carbohydrate Metabolism↗