Predicting tropical plant physiology from leaf and canopy spectroscopy

A broad regional understanding of tropical forest leaf photosynthesis has long been a goal for tropical forest ecologists, but it has remained elusive due to difficult canopy access and high species diversity. Here we develop an empirical model to predict sunlit, light-saturated, tropical leaf photosynthesis using leaf and simulated canopy spectra. To develop this model, we used partial least squares (PLS) analysis on three tropical forest datasets (159 species), two in Hawaii and one at the biosphere 2 laboratory (B2L). For each species, we measured light-saturated photosynthesis (A), light and CO₂ saturated photosynthesis (A _max), respiration (R), leaf transmittance and reflectance spectra (400–2,500 nm), leaf nitrogen, chlorophyll a and b, carotenoids, and leaf mass per area (LMA). The model best predicted A [r ²  = 0.74, root mean square error (RMSE) = 2.9 μmol m⁻² s⁻¹)] followed by R (r ²  = 0.48), and A _max (r ²  = 0.47). We combined leaf reflectance and transmittance with a canopy radiative transfer model to simulate top-of-canopy reflectance and found that canopy spectra are a better predictor of A (RMSE = 2.5 ± 0.07 μmol m⁻² s⁻¹) than are leaf spectra. The results indicate the potential for this technique to be used with high-fidelity imaging spectrometers to remotely sense tropical forest canopy photosynthesis.     

Intra- and inter-specific variation in canopy photosynthesis in a mixed deciduous forest

 Within the same forest, photosynthesis can vary greatly among species and within an individual tree. Quantifying the magnitude of variation in leaf-level photosynthesis in a forest canopy will improve our understanding of and ability to model forest carbon cycling. This information requires extensive sampling of photosynthesis in the canopy. We used a 22-m-tall, four-wheel-drive aerial lift to reach five to ten leaves from the tops of numerous individuals of several species of temperate deciduous trees in central Massachusetts. The goals of this study were to measure light-saturated photosynthesis in co-occurring canopy tree species under field conditions, and to identify sampling schemes appropriate for canopy tree studies with challenging logistics. Photosynthesis differed significantly among species. Even though all leaves measured were canopy-top, sun-acclimated foliage, the more shade-tolerant species tended to have lower light-saturated photosynthetic rates (P _max) than the shade-intolerant species. Likewise, leaf mass per area (LMA) and nitrogen content (N) varied significantly between species. With only one exception, the shade-tolerant species tended to have lower nitrogen content on an area basis than the intolerant species, although the LMA did not differ systematically between these ecological types. Light-saturated P _max rates and nitrogen content, both calculated on either an area or a mass basis, and the leaf mass to area ratio, significantly differed not only among species, but also among individuals within species (P<0.0001 for both). Differences among species accounted for a greater proportion of variance in the P _max rates and the nitrogen content than the differences among individuals within a species (58.5–78.8% of the total variance for the measured parameters was attributed to species-level differences versus 5.5–17.4% of the variance was attributed to differences between individual trees of a given species). Furthermore, more variation is accounted for by differences among leaves in a single individual tree, than by differences among individual trees of a given species (10.7–30.4% versus 5.5–17.4%). This result allows us to compare species-level photosynthesis, even if the sample size of the number of trees is low. This is important because studies of canopy-level photosynthesis are often limited by the difficulty of canopy access. As an alternative to direct canopy access measurements of photosynthesis, it would be useful to find an ”easy-to-measure” proxy for light-saturated photosynthetic rates to facilitate modeling forest carbon cycling. Across all species in this study, the strongest correlation was between nitrogen content expressed on an area basis (mmol m^–2, N _area) and light-saturated P _max rate (μmol m^–2 s^–1, P _maxarea) (r ²=0.511). However, within a given species, leaf nitrogen was not tightly correlated with photosynthesis. Our sampling design minimized intra-specific leaf-level variation (i.e., leaves were taken only from the top of the canopy and at only one point in the season). This implies that easy-to-measure trends in nitrogen content of leaves may be used to predict the species-specific light-saturated P _max rates. Received: 16 March 1996 / Accepted: 16 August 1996     

Reconciling the apparent difference between mass- and area-based expressions of the photosynthesis-nitrogen relationship

The relationship between photosynthetic carbon assimilation (A _max) and leaf nitrogen content (N _leaf) can be expressed on either a leaf area basis (A _area vs N _area) or a leaf mass basis (A _mass vs N _mass). Dimensional analysis shows that the units for the slope of this relationship are the same for both expressions (μmol [CO₂] g⁻¹ [N] s⁻¹). Thus the slope measures the change in CO₂ assimilation per gram of nitrogen, independent of leaf mass or leaf area. Although they have the same units, large differences between the area and mass-based slopes have been observed over a broad range of taxonomically diverse species. Some authors have claimed that regardless of these differences, the fundamental nature of the A _max-N _leaf relationship is independent of the units of expression. In contrast, other authors have claimed that the area-based A _max-N _leaf relationship is fundamentally different from the mass-based relationship because of interactions between A _max, N _leaf, and leaf mass per area (LMA, g [leaf] m⁻² [leaf]). In this study we consider the mathematical relationships involved in the transformation from mass- to area-based expressions (and vice versa), and the implications this transformation has for the slope of the A _max-N _leaf relationship. We then show that the slope of the relationship is independent of the units of expression when the effect of LMA is controlled statistically using a multiple regression. The validity of this hypothesis is demonstrated using 13 taxonomically and functionally diverse C₃ species. This analysis shows that the slope of the A _max-N _leaf relationship is similar for the mass- and area-based expressions and that significant errors in the estimate of the slope can arise when the effect of LMA is not controlled. Received: 7 May 1998 / Accepted: 19 October 1998     

Leaf gas exchange of trees in old-growth and young secondary forest stands in Sulawesi, Indonesia

In the tropics, old-growth forests are converted to other land cover types at a high rate and young secondary forest may gain in importance. Information on associated changes in leaf gas exchange and other leaf traits can be valuable for modelling biogeochemical fluxes under altered land-use patterns. We studied in situ photosynthetic parameters and stomatal conductance for water vapour in eight abundant tree species of young secondary forest and eight tree species of natural old-growth forest in Central Sulawesi, Indonesia. In sun leaves, the average maximal stomatal conductance (g _smax) in the secondary forest (SF) species was 2.1 times higher than in the old-growth forest (OGF) species. Species with a high g _smax reduced g _s sharply when vapour pressure deficit of the air increased, whereas species with a low g _smax were much less sensitive to air humidity. For area-based photosynthetic capacity (A _max-area), the SF species had a 2.3 times higher average than the OGF species. For both, g _smax and A _max-area the variation among species was higher in the OGF than in the SF. When all tree species (n=16) are considered, species means of specific leaf area (SLA), leaf N concentration and leaf P concentration were significantly correlated with g _smax and A _max-area. The strong correlation between A _max-area and foliar P (r ²=0.8) is remarkable as the alluvial soils in the study region are rich in nutrients. If the eight OGF species are analysed separately, the only significant correlation was observed between SLA and mass-based A _max; in the SF species strong correlations were found between leaf size and A _max-area and g _smax. These results show that the conversion of old-growth forest to young secondary forest in Sulawesi significantly alters tree leaf gas exchange characteristics and that chemical and structural leaf traits can be used for the prediction of these changes. The best correlations between leaf gas exchange parameters and leaf traits were obtained by different traits in the SF species, the OGF species and the entire pool of studied species.     

Changes in needle nitrogen partitioning and photosynthesis during 80 years of tree ontogeny in Picea abies

Needle nitrogen partitioning and photosynthesis of Norway spruce were studied in a forest chronosequence in Järvselja Experimental Forest, Estonia. Current- and previous-year shoots were sampled from upper and lower canopy positions in four stands, ranging in age from 13 to 82 years. A/c _i curves were determined to obtain maximum carboxylation rate (V _cmax) and maximum rate of electron transport (J _max), whereas needle nitrogen partitioning into carboxylation (P _R), bioenergetics associated with electron transport (P _B) and thylakoid light harvesting components (P _L) was calculated from the values of V _cmax, J _max and leaf chlorophyll concentration. The greatest changes in studied needle characteristics took place between tree ages of 13 and 26 years, and this pattern was independent of needle age and canopy position. Needle mass per projected area (LMA) was lowest in the 13-year-old stand and mass-based nitrogen concentration (N^M) was generally highest in that stand. The values of LMA were significantly higher and those of N^M lower in the 26-year-old stand. Mass-based V _cmax and J _max were highest in the 13-year-old stand. Area-based photosynthetic capacity was independent of tree age. The proportion of photosynthetic nitrogen (P _R, P _B and P _L) was highest and that of non-photosynthetic nitrogen lowest in the 13-year-old stand. Current-year needles had lower LMA and P _L, but higher photosynthetic capacity compared to 1-year-old foliage. Needles from lower canopy positions exhibited lower LMA, area-based nitrogen concentration and photosynthetic capacity than needles from upper canopy. The period of substantial reductions in needle photosynthetic capacity and changes in nitrogen partitioning coincides with the onset of reproductive phase during tree ontogeny.     

Photosynthetic plasticity of two rain forest shrubs across natural gap transects

Summary Photosynthetic plasticity of two congeneric shrub species growing under natural field conditions was compared along transects spanning two canopy gaps in a Costa Rican rain forest. Piper arieianum is a shadetolerant species common in successional and mature forests, whereas P. sancti-felicis is a pioneer species abundant in abandoned clearings and large gaps. Twenty potted cuttings of each species were placed at regular intervals along two east-west transects crossing a small branch-fall gap and a large tree-fall gap. Along the transects, the percent of full sun photon flux density varied from less than 2% to 45%. After six months of growth under these conditions, leaves were monitored for incident photon flux density, photographic measures of light availability, photosynthetic capacity (A_max), leaf nitrogen content, leaf chlorophyll content, and specific leaf mass. Although both species demonstrated considerable plasticity in A_max across gap transects, P. sancti-felicis leaves had a superior capacity to track closely variation in light availability, particularly in the larger gap. For regressions of A_max on measures of light availability, P. sancti-felicis consistently showed a 3.5 to 5-fold higher coefficient of determination (R²) and a 3 to 4-fold higher slope than P. arieianum. In both species leaf nitrogen content per leaf area increased significantly with light availability, although P. sancti-felicis, again, showed a much stronger relationship between these variables. Across the transects, mean chlorophyll content per unit leaf area did not differ significantly between the species, whereas mean chlorophyll content per unit leaf dry mass was 3-times greater in leaves of P. sancti-felicis. Piper arieianum exhibited highly significant increases in chlorophyll a:b ratio with increased light availability, whereas P. sancti-felicis lacked significant variation in this trait across a gradient of light availability. Mean specific leaf mass did not vary significantly between species across the gap transects. The nature of the light acclimatory response differs quantitatively and qualitatively between these species. An important constraint on light acclimation of the shade-tolerant P. arieianum is its inability to increase photosynthetic nitrogen-use efficiency under conditions of high light availability. The lack of plasticity in chlorophyll a:b ratios does not restrict light acclimation of A_max in P. sancti-felicis. Leaves of P. arieianum exhibited symptoms of chronic photoinhibition in exposed microsites within the large gap. Species differences in the capacity to finely adjust A_max across a wide range of light conditions may be attributed to their maximum growth potential. Light acclimation in species with low maximum growth potential may be constrained at the cellular level by rates of protein and chlorophyll synthesis and at the whole-plant level by low maximum rates of uptake and supply of nutrients and water. For P. arieianum, restriction of photosynthetic plasticity is likely to limit competitive abilities of plants in high-light conditions of large gaps and clearings, whereas observed habitat restrictions for P. sancti-felicis do not appear to depend upon the highly-developed capacity for adjustment of A_max observed in this species.     

Contrasting leaf characteristics of trees and lianas in secondary and mature forests in southwestern China

We compared variation in sun-canopy leaf anatomy, morphology and photosynthetic rates of coexisting woody species (trees and lianas) in an 8-year-old secondary forest (SF) and mature forest (MF) in the wet season in Xishuangbanna, SW China. Variability of leaf traits of 66 species within growth-form groups in each forest was quantified using coefficients of variation (CV). For the mean values, the woody species in the SF had significantly higher leaf thickness and stomatal density, but lower nonmesophyll/mesophyll ratios than those in the MF. The average leaf area and leaf mass area (LMA) in the studied woody species did not change greatly during the successional process, but differed significantly between the growth forms, with trees having higher values than lianas. The light-saturated photosynthetic rate per unit leaf area (A _a) of the woody species in the SF ranged from 11.2 to 34.5 μmol m⁻² s⁻¹, similarly to pioneer tree species from literature data in southeast Asia. The A _a and photosynthetic nitrogen-use efficiency (PNUE) were significantly higher than those in the MF; whereas A _a in the MF ranged between 9 to 21 μmol m⁻² s⁻¹, with similar values between lianas and trees. For all woody species in both SF and MF, there were no significant differences in the average values of the CV of all measured variables for both lianas and trees. However, considerable variation in leaf anatomy, morphology, and photosynthetic rates within both growth forms and forests existed, as well as substantial variation in leaf size and stomatal density. We concluded that the tropical woody species formed a heterogeneous functional group in terms of leaf morphology and physiology in both secondary and mature forests.     

Photosynthetic parameters,dark respiration and leaf traits in the canopy of a Peruvian tropical montane cloud forest

Few data are available describing the photosynthetic parameters of the leaves of tropical montane cloud forests (TMCF). Here, we present a study of photosynthetic leaf traits (V _cmax and J _max), foliar dark respiration (R _d), foliar nitrogen (N) and phosphorus (P), and leaf mass per area (LMA) throughout the canopy for five different TMCF species at 3025 m a.s.l. in Andean Peru. All leaf traits showed a significant relationship with canopy height when expressed on an area basis, and V _cmax-area and J _max-area almost halved when descending through the TMCF canopy. When corrected to a common temperature, average V _cmax and J _max on a leaf area basis were similar to lowland tropical values, but lower when expressed on a mass basis, because of the higher TMCF LMA values. By contrast, R _d on an area basis was higher than found in tropical lowland forests at a common temperature, and similar to lowland forests on a mass basis. The TMCF J _max–V _cmax relationship was steeper than in other tropical biomes, and we propose that this can be explained by either the light conditions or the relatively low VPD in the studied TMCF. Furthermore, V _cmax had a significant—though relatively weak and shallow—relationship with N on an area basis, but not with P, which is consistent with the general hypothesis that TMCFs are N rather than P limited. Finally, the observed V _cmax–N relationship (i.e., maximum photosynthetic nitrogen use efficiency) was distinctly different from those in tropical and temperate regions, probably because the TMCF leaves compensate for reduced Rubisco activity in cool environments.     

Differential leaf traits of a neotropical tree Cariniana legalis (Mart.) Kuntze (Lecythidaceae): comparing saplings and emergent trees

Cariniana legalis is an emergent tree that reaches the upper canopy in Brazilian Semideciduous Forest. Spatial contrasts in microclimatic conditions between the upper canopy and understorey in a forest may affect morpho-physiological leaf traits. In order to test the hypothesis that the upper canopy is more stressful to leaves than a gap environment we compared emergent trees of C. legalis, 28–29 m in height to gap saplings, 6–9 m in height, for the following parameters: leaf area, leaf mass area (LMA the dry weight:leaf area ratio), leaf thickness, leaf anatomical parameters, stomata conductance, and chlorophyll a fluorescence. Leaves from emergent trees had smaller leaf areas but greater LMA compared to saplings. Leaf thickness, palisade layer thickness, and stomatal density were higher for emergent trees than for saplings. The opposite pattern was observed for spongy layer thickness and spongy/palisade ratio. Stomatal conductance was also higher for emergent tree leaves than for sapling leaves, but the magnitude of depression on stomatal conductance near midday was more pronounced in emergent trees. The potential quantum yield of photosystem II, as determined by the F _v/F _m ratio was lower for leaves from saplings. The lower values of stomatal conductance, indicating restriction in CO₂ diffusion into the mesophyll can be related to higher photoinhibition observed in the saplings. Leaves from emergent trees and saplings exhibited similar values for apparent electron transport rates and non-photochemical quenching. Our results suggest that changes in leaf traits could be associated to dry conditions at the upper canopy as well as to the ontogenetic transition between sapling/emergent tree life stages.     

Remote sensing of forest gas exchange: Considerations derived from a tomographic perspective

The global exchange of gas (CO₂, H₂O) and energy (sensible and latent heat) between forest ecosystems and the atmosphere is often assessed using remote sensing (RS) products. Although these products are essential in quantifying the spatial variability of forest–atmosphere exchanges, large uncertainties remain from a measurement bias towards top of canopy fluxes since optical RS data are not sensitive for the vertically integrated forest canopy. We hypothesize that a tomographic perspective opens new pathways to advance upscaling gas exchange processes from leaf to forest stands and larger scales. We suggest a 3D modelling environment comprising principles of ecohydrology and radiative transfer modelling with measurements of micrometeorological variables, leaf optical properties and forest structure, and assess 3D fields of net CO₂ assimilation (A_n) and transpiration (T) in a Swiss temperate forest canopy. 3D simulations were used to quantify uncertainties in gas exchange estimates inherent to RS approaches and model assumptions (i.e. a big‐leaf approximation in modelling approaches). Our results reveal substantial 3D heterogeneity of forest gas exchange with top of canopy A_n and T being reduced by up to 98% at the bottom of the canopy. We show that a simplified use of RS causes uncertainties in estimated vertical gas exchange of up to 300% and that the spatial variation of gas exchange in the footprint of flux towers can exceed diurnal dynamics. We also demonstrate that big‐leaf assumptions can cause uncertainties up to a factor of 10 for estimates of A_n and T. Concluding, we acknowledge the large potential of 3D assessments of gas exchange to unravelling the role of vertical variability and canopy structure in regulating forest–atmosphere gas and energy exchange. Such information allows to systematically link canopy with global scale controls on forest functioning and eventually enables advanced understanding of forest responses to environmental change.     

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content

Background and Aims

Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species.

Methods

Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0·5 Hz over 9–17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N_area), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A_max), dark respiration (R_d), apparent quantum yield (ϕ) and curvature (θ)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A_daily) for each leaf.

Key Results

Leaf N_area and A_max changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A_daily did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species.

Conclusions

The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple ‘big-leaves’ in which all constituent ‘small leaves’ use PPFD with the same photosynthetic efficiency.Key words: Optimal resource allocation, nitrogen, photosynthetic capacity, leaf mass per area, tropical trees, radiation use efficiency, scaling, leaf angle, canopy architecture, big leaf model     

Irradiance acclimation,capture ability,and efficiency in invasive and non-invasive alien plant species

We tested the hypothesis that invasive (IN) species could capture resources more rapidly and efficiently than noninvasive (NIN) species. Two IN alien species, Ageratina adenophora and Chromolaena odorata, and one NIN alien species, Gynura sp. were compared at five irradiances. Photon-saturated photosynthetic rate (P _max), leaf mass (LMA) and nitrogen content (N_A) per unit area, and photosynthetic nitrogen utilization efficiency (PNUE) increased significantly with irradiance. LMA, N_A, and PNUE all contributed to the increased P _max, indicating that both morphological and physiological acclimation were important for the three alien species. Under stronger irradiance, PNUE was improved through changes in N allocation. With the increase of irradiance, the amount of N converted into carboxylation and bioenergetics increased, whereas that allocated to light-harvesting components decreased. The three alien species could adequately acclimate to high irradiance by increasing the ability to utilize and dissipate photon energy and decreasing the efficiency of photon capture. The two IN species survived at 4.5 % irradiance while the NIN species Gynura died, representing their different invasiveness. Ageratina generally exhibited higher respiration rate (R _D) and N_A. However, distinctly higher P _max, PNUE, P _max/R _D, or P _max/LMA were not detected in the two invasive species, nor was lower LMA. Hence the abilities to capture and utilize resources were not always associated with invasiveness of the alien species.     

An In Situ Leaf and Branch Warming Experiment in the Amazon

Leaves and branches of mature trees, lianas, and gap species were warmed in an Amazonian forest for 4 mo to observe the effect of warming on photosynthesis, stomatal conductance, and transpiration. Electric resistance heaters increased air temperatures near the leaves by approximately 2°C. Sunlit leaf temperatures increased by 2–3°C on average, but during some periods leaf temperatures increased by >5°C. Maximum photosynthesis (A_max) decreased significantly in the warmed leaves vs. the control leaves over the 13‐wk study period with an average decrease in A_max of 1.4 μmol/m²s (19% decrease from a mean A_max of 7.2 μmol/m²s) when measured at 30°C and there were no signs of acclimation to higher temperatures within existing leaves. The decline in A_max was likely due to irreversible temperature damage caused by very high leaf temperatures and not due to C_i limitation of carboxylation. Warming had a larger negative impact on A_max in canopy level tree species than other tested functional groups such as lianas or gap species. Transpiration did not significantly increase in the warmed leaves compared with the control group. This study indicates that increased temperatures due to global warming could potentially decrease future tropical forest carbon uptake by a significant amount. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

Leaf Longevity,Construction, and Maintenance Costs of Three Mangrove Species Under Field Conditions

This study assessed the effect of leaf age on construction cost (CC) in the mangrove species Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle growing in their natural habitat. Leaf osmolality values were species-specific, the highest in A. germinans (1 693 mmol kg^–1) and the lowest in L. racemosa (1 270 mmol kg^–1). In the three species, contents of chlorophyll (a+b) (Chl_a+b) and nitrogen (N) per unit of leaf area were maximal in adult leaves and tended to decline with age. Leaf mass to leaf area ratio (LMA) and ash content increased during leaf ageing. Similarly, as leaves aged, a significant increase in leaf construction cost per leaf area (CC_a) was observed, while per leaf mass (CC_m) it remained almost constant, suggesting a sustained production of leaf compounds as leaves became older. CC was positively correlated with LMA and heat of combustion (Hc) per leaf area, suggesting differences among species in the quantity and composition of expensive compounds. Leaf half lifetime (t_0.5) showed contrasting values in the three mangrove species (60, 111, and 160 d in L. racemosa, R. mangle, and A. germinans, respectively). Overall, L. racemosa was the species with less expensive leaves to construct while leaves of A. germinans and R. mangle had the highest CC_m and CC_a, respectively. Leaf longevity was positively correlated with the ratio between CC and maximum photosynthetic rate (P _max), clearly showing the existence of a balance between leaf costs and benefits.     

Relationships of leaf dark respiration to leaf nitrogen, specific leaf area and leaf life-span: a test across biomes and functional groups

Based on prior evidence of coordinated multiple leaf trait scaling, we hypothesized that variation among species in leaf dark respiration rate (R _d) should scale with variation in traits such as leaf nitrogen (N), leaf life-span, specific leaf area (SLA), and net photosynthetic capacity (A _max). However, it is not known whether such scaling, if it exists, is similar among disparate biomes and plant functional types. We tested this idea by examining the interspecific relationships between R _d measured at a standard temperature and leaf life-span, N, SLA and A _max for 69 species from four functional groups (forbs, broad-leafed trees and shrubs, and needle-leafed conifers) in six biomes traversing the Americas: alpine tundra/subalpine forest, Colorado; cold temperate forest/grassland, Wisconsin; cool temperate forest, North Carolina; desert/shrubland, New Mexico; subtropical forest, South Carolina; and tropical rain forest, Amazonas, Venezuela. Area-based R _d was positively related to area-based leaf N within functional groups and for all species pooled, but not when comparing among species within any site. At all sites, mass-based R _d (R _d-mass) decreased sharply with increasing leaf life-span and was positively related to SLA and mass-based A _max and leaf N (leaf N _mass). These intra-biome relationships were similar in shape and slope among sites, where in each case we compared species belonging to different plant functional groups. Significant R _d-mass−N _mass relationships were observed in all functional groups (pooled across sites), but the relationships differed, with higher R _d at any given leaf N in functional groups (such as forbs) with higher SLA and shorter leaf life-span. Regardless of biome or functional group, R _d-mass was well predicted by all combinations of leaf life-span, N _mass and/or SLA (r ²≥ 0.79, P < 0.0001). At any given SLA, R _d-mass rises with increasing N _mass and/or decreasing leaf life-span; and at any level of N _mass, R _d-mass rises with increasing SLA and/or decreasing leaf life-span. The relationships between R _d and leaf traits observed in this study support the idea of a global set of predictable interrelationships between key leaf morphological, chemical and metabolic traits. Received: 23 May 1997 / Accepted: 16 December 1997     

Photosynthetic Characteristics,Dark Respiration,and Leaf Mass Per Unit Area in Seedlings of Four Tropical Tree Species Grown Under Three Irradiances

We investigated the effect of growth irradiance (I) on photon-saturated photosynthetic rate (P _max), dark respiration rate (R _D), carboxylation efficiency (CE), and leaf mass per unit area (LMA) in seedlings of the following four tropical tree species with contrasting shade-tolerance. Anthocephalus chinensis (Rubiaceae) and Linociera insignis (Oleaceae) are light-demanding, Barringtonia macrostachya (Lecythidaceae) and Calophyllum polyanthum (Clusiaceae) are shade-tolerant. Their seedlings were pot-planted under shading nets with 8, 25, and 50 % daylight for five months. With increase of I, all species displayed the trends of increases of LMA, photosynthetic saturation irradiance, and chlorophyll-based P _max, and decreases of chlorophyll (Chl) content on both area and mass bases, and mass-based P _max, R _D, and CE. The area-based P _max and CE increased with I for the light-demanders only. Three of the four species significantly increased Chl-based CE with I. This indicated the increase of nitrogen (N) allocation to carboxylation enzyme relative to Chl with I. Compared to the two shade-tolerants, under the same I, the two light-demanders had greater area- and Chl-based P _max, photosynthetic saturation irradiance, lower Chl content per unit area, and greater plasticity in LMA and area- or Chl-based P _max. Our results support the hypothesis that light-demanding species is more plastic in leaf morphology and physiology than shade-tolerant species, and acclimation to I of tropical seedlings is more associated with leaf morphological adjustment relative to physiology. Leaf nitrogen partitioning between photosynthetic enzymes and Chl also play a role in the acclimation to I.     

温带针阔混交林叶片性状随树冠垂直高度的变化规律

自然界中,森林植物叶片的生长随树冠高度呈现明显的垂直分布现象;然而,有关叶片性状随着树冠垂直高度增加的变化规律仍不清楚。为了更好地揭示植物叶片对光环境变化的适应策略以及对资源的利用能力,有必要深入探讨叶片性状与冠层高度的定量关系及其内在调控机制。以中国广泛分布的温带针阔混交林为对象,选取8种主要树种为研究对象(白桦、蒙古栎、水曲柳、大青杨、色木槭、千金榆、核桃楸和红松),通过测定这些物种9个冠层高度的叶片比叶面积(SLA)、叶片干物质含量(LDMC)、叶片氮含量(N)、叶片磷含量(P)、氮磷比(N∶P)和叶绿素含量(Chl)等属性,探讨了针阔混交林叶片性状的差异以及各性状之间的相关关系,进而揭示叶片性状随树冠垂直高度的变化规律。实验结果表明:1)温带针阔混交林内优势树种的部分叶片性状在不同冠层高度之间差异显著。2)随着树冠垂直高度的增加,SLA、LDMC、N、P、N∶P和Chl呈现不同的变化趋势。其中,阔叶树种SLA随着树冠垂直高度的增加而减小;所有树种的LDMC随着树冠垂直高度的增加而增加;不同树种的N、P、N∶P和Chl随着树冠垂直高度的变化规律存在差异。3)对于温带针阔混交林冠层中,SLA与N、P、N∶P均存在显著的正相关关系,高SLA伴随着高的N、P、N∶P,表明植物通过SLA与N、P等性状的协同来提高叶片的光合作用(或对光热资源的利用效率)。本研究通过定量分析探讨温带针阔混交林叶片性状随冠层高度的变化规律,一定程度地揭示了树木对光、热和水资源竞争的适应机制,以及植物叶片的资源利用和分配策略,不仅拓展了传统性状研究的范畴,其相关研究结论也有助于树木生长模型的构建和优化。     

Quantitative trait loci controlling water use efficiency and related traits in Quercus robur L.

Genetic variation for intrinsic water use efficiency (W _i) and related traits was estimated in a full-sib family of Quercus robur L. over 3 years. The genetic linkage map available for this F1 family was used to locate quantitative trait loci (QTL) for W _i, as estimated by leaf carbon stable isotope composition (δ ¹³C) or the ratio of net CO₂ assimilation rate (A) to stomatal conductance to water vapour (g _w) and related leaf traits. Gas exchange measurements were used to standardize estimates of A and g _w and to model the sensitivity of g _w to leaf-to-air vapour pressure deficit (sg^VPD). δ ¹³C varied by more than 3‰ among the siblings, which is equivalent to 40% variation of W _i. Most of the studied traits exhibited high clonal mean repeatabilities (>50%; proportion of clonal mean variability in global variance). Repeatabilities for δ ¹³C, leaf mass per area (LMA) and leaf nitrogen content were higher than 70%. For δ ¹³C, ten QTLs were detected, one of which was detected repeatedly for all 3 years and consistently explained more than 20% of measured variance. Four genomic regions were found in which co-localizing traits linked variation in W _i to variations in leaf chlorophyll and nitrogen content, LMA and sg^VPD. A positive correlation using clonal means between δ ¹³C and A/g _w, as well as a co-localisation of QTL detected for both traits, can be seen as validation of the theoretical model linking the genetic architecture of these two traits.     

Leaf traits and photosynthetic parameters of saplings and adult trees of co-existing species in a temperate broad-leaved forest

In Central European forestry the establishment of broad-leaved mixed forests is attaining increasing importance, but little information exists about gas exchange characteristics of some of the tree species involved, which are less abundant today. In an old-growth forest in Central Germany (Hainich, Thuringia), (i) I compared morphological and chemical leaf traits that are indicative of leaf gas exchange characteristics among eight co-existing species, and (ii) analysed photosynthetic parameters of saplings and adult trees (lower and upper canopy level) in four of these species (Acer pseudoplatanus L., Carpinus betulus L., Fraxinus excelsior L. and Tilia platyphyllos Scop.).Leaves from the upper canopy in the eight species studied varied significantly in their specific leaf area (12.9–19.4 m² kg⁻¹), stomatal density (125–313 stomata mm⁻²), leaf nitrogen concentration (95–157 mmol N m⁻²) and δ¹³C content (–27.81 to –25.85‰). F. excelsior and C. betulus were largely contrasting species, which suggests that the species, which were studied in more detail, include the widest difference in leaf gas exchange among the co-existing species. The saplings of the four selected species exhibited shade acclimated leaves with net photosynthesis rates at saturating irradiance (A_max) between 5.0 and 6.4 μmol m⁻² s⁻¹. In adult trees A_max of fully sunlit leaves was more variable and ranged from 10.5 (C. betulus) to 16.3 μmol m⁻² s⁻¹ (F. excelsior). However, less negative δ¹³C values in F. excelsior sun leaves point to a strong limitation in gas exchange. In the lower canopy of adult trees A_max of F. excelsior (12.0 μmol m⁻² s⁻¹) was also greater than that of A. pseudoplatanus, C. betulus and T. platyphyllos (5.0–5.6 μmol m⁻² s⁻¹). This can be explained by the small leaf area and the absence of shade leaves in mature F. excelsior trees. Thus, a considerable variation in leaf traits and gas exchange was found among the co-existing tree species. The results suggest that species-specific characteristics increase the spatial heterogeneity of canopy gas exchange and should be taken into account in the interpretation and prediction of gas flux from mixed stands.In der Forstwirtschaft Mitteleuropas gewinnt die Begründung von Laubmischwäldern zunehmend an Bedeutung, aber über Eigenschaften im Gasaustausch einiger beteiligter Baumarten, die heute nicht so häufig sind, ist wenig bekannt. In einem Altbestand in Mitteldeutschland (Hainich, Thüringen) habe ich (i) morphologische und chemische Eigenschaften von Sonnenblättern, die Hinweise auf Charakteristika im Blattgaswechsel geben, an acht koexistierenden Baumarten untersucht, und (ii) Photosyntheseparameter von juvenilen und adulten Bäumen (unteres und oberes Kronenniveau) von vier dieser Arten (Acer pseudoplatanus L., Carpinus betulus L., Fraxinus excelsior L. and Tilia platyphyllos Scop.) erhoben.Blätter aus dem oberen Kronenraum der acht untersuchten Arten variierten signifikant in der spezifischen Blattfläche (12.9–19.4 m² kg⁻¹), der Stomatadichte (125–313 Stomata mm⁻²), dem Blattstickstoffgehalt (95–157 mmol N m⁻²) und den δ¹³C-Werten (–27.81 bis –25.85‰). In diesem Kollektiv zeigten F. excelsior und C. betulus groβe Unterschiede, was darauf hindeutet, dass die Arten, die genauer untersucht wurden, die Spannweite an Gaswechseleigenschaften unter den koexistierenden Baumarten umfassen. Die Jungpflanzen der vier ausgewählten Arten besaßen Schattenblätter, deren Netto-Photosyntheserate bei hoher Lichtintensität (A_max) zwischen 5.0 and 6.4 μmol m⁻² s⁻¹ variierte. An Sonnenblättern von Altbäumen war A_max variabler und lag zwischen 10.5 (C. betulus) und 16.3 μmol m⁻² s⁻¹ (F. excelsior). Allerdings weisen hohe δ¹³C-Werte in Sonnenblättern von F. excelsior auf eine starke Limitierung des Gasaustauschs hin. Auch in der unteren Krone der Altbäume war A_max von F. excelsior (12.0 μmol m⁻² s⁻¹) höher als A_max von A. pseudoplatanus, C. betulus und T. platyphyllos (5.0–5.6 μmol m⁻² s⁻¹). Dies kann durch die geringe Blattfläche und die Abwesenheit von Schattenblättern in der Krone adulter Bäume von F. excelsior erklärt werden. Zwischen den koexistierenden Baumarten wurde somit in Bezug auf Blatteigenschaften und Photosyntheseparameter eine erhebliche Variation festgestellt. Die Ergebnisse legen nahe, dass artspezifische Eigenschaften die räumliche Heterogenität des Gaswechsels im Kronenraum erhöhen und bei der Interpretation und Vorhersage von Gasflüssen über Mischbeständen berücksichtigt werden sollten.     

Effects of leaf position and nitrogen supply on the expansion of leaves of field grown sunflower (Helianthus annuus L.)

Leaf growth responses to N supply and leaf position were studied using widely-spaced sunflower plants growing under field conditions. Both N supply (range 0.25 to 11.25 g added N per plant) and leaf position significantly (p=0.001) affected maximum leaf area (LA_max) of target leaves through variations in leaf expansion rate (LER); effects on duration of expansion were small. Specific leaf nitrogen (SLN, g N m^-2) fell quite rapidly during the initial leaf expansion phase (LA < 35% LA_max) but leveled off during the final 65% increase of leaf area. This pattern held across leaf positions and N supply levels. Leaf nitrogen accumulation after 35% LA_max continued up to achievement of LA_max; reductions in the higher SLN characteristic of the initial phase were insufficient to cover the nitrogen requirements for expansion during the final phase. LER in the quasi-linear expansion phase (35 to 100% of LA_max) was strongly associated with SLN above a threshold that varied with leaf position (mean 1.79±0.225 g N m^-2). This contrasts with the response of photosynthesis at high irradiance to SLN, which has previously been shown to have a threshold of 0.3 g N m^-2; in the present work saturation of photosynthetic rate was evident when SLN reached 1.97 g N m^-2. Thus, once the area of a leaf exceeds 35% of LA_max, expansion proceeds provided SLN values are close to the levels required for maximum photosynthesis. However, growth of leaves during the initial expansion phase ensures a minimum production of leaf area even at low N supply levels.