Co-optimal distribution of leaf nitrogen and hydraulic conductance in plant canopies

Leaf properties vary significantly within plant canopies, due to the strong gradient in light availability through the canopy, and the need for plants to use resources efficiently. At high light, photosynthesis is maximized when leaves have a high nitrogen content and water supply, whereas at low light leaves have a lower requirement for both nitrogen and water. Studies of the distribution of leaf nitrogen (N) within canopies have shown that, if water supply is ignored, the optimal distribution is that where N is proportional to light, but that the gradient of N in real canopies is shallower than the optimal distribution. We extend this work by considering the optimal co-allocation of nitrogen and water supply within plant canopies. We developed a simple 'toy' two-leaf canopy model and optimized the distribution of N and hydraulic conductance (K) between the two leaves. We asked whether hydraulic constraints to water supply can explain shallow N gradients in canopies. We found that the optimal N distribution within plant canopies is proportional to the light distribution only if hydraulic conductance, K, is also optimally distributed. The optimal distribution of K is that where K and N are both proportional to incident light, such that optimal K is highest to the upper canopy. If the plant is constrained in its ability to construct higher K to sun-exposed leaves, the optimal N distribution does not follow the gradient in light within canopies, but instead follows a shallower gradient. We therefore hypothesize that measured deviations from the predicted optimal distribution of N could be explained by constraints on the distribution of K within canopies. Further empirical research is required on the extent to which plants can construct optimal K distributions, and whether shallow within-canopy N distributions can be explained by sub-optimal K distributions.     

Seasonal changes in leaf nitrogen pools in two Salix species

Leaf nitrogen distribution pattern was studied four times during the growing season in a 2-year-old Salix viminalis L. and Salix dasyclados Wimm. plantation in Estonia. We measured the vertical distributions of leaf nitrogen concentration, dry mass, leaf area and light environment (as fractional transmission of diffuse irradiance, a(d)) in the canopy. The light-independent nitrogen pool was evaluated as the intercept of the leaf nitrogen concentration versus a(d) relationship, and the nondegradable nitrogen pool was evaluated as the nitrogen remaining in abscised leaves. A strong vertical gradient of mass-based leaf nitrogen concentration was detected at the beginning of the growing season, and decreased steadily during canopy development. This decline had at least three causes: (1) the amount of nitrogen in the foliage was larger at the beginning of the growing season than at the end of the growing season, probably because of pre-existing root systems; (2) with increasing leaf area index (LAI) during the growing season, the proportion of leaf nitrogen in total canopy nitrogen that could be redistributed (light-dependent nitrogen pool) decreased; and (3) the photosynthetic photon flux density gradient inside the canopy changed during the season, most probably because of changes in leaf area and leaf angle distributions. Total canopy nitrogen increased almost proportionally to LAI, whereas the light-dependent nitrogen pool had a maximum in August. Also, the proportion of the light-dependent nitrogen pool in the total canopy nitrogen decreased steadily from 65.2% in June to 17.2% in September in S. dasyclados and from 63.3 to 15.1% in S. viminalis. The degradable nitrogen pool was always bigger than the light-dependent nitrogen pool.     

Diurnal variation in leaf extension of Salix viminalis at two nitrogen supply rates

To investigate how nitrogen supply might affect the biophysical factors controlling diurnal variation in leaf extension, pot-grown Salix viminalis L. were supplied with nitrogen at a low relative addition rate of 0.05 g N g(-1) N day(-1) (low N) or were given free access to all nutrients (high N). Leaf extension, turgor pressure, turgor after stress relaxation and the plastic extensibility of leaf tissue were determined for growing leaves every 4 h during two days of clear skies in August. Plants in the high-N treatment had a significantly higher relative growth rate, dry weight, shoot/root ratio, leaf nitrogen concentration, total leaf area, final area of single leaves and epidermal cell size than plants in the low-N treatment. The periodicity of leaf extension was similar in both treatments with high values during the afternoon and early evening, and negligible values during the night and in the early morning. The maximum rate of leaf extension was higher in high-N than in low-N plants. Leaf water potential and leaf osmotic potential decreased in the morning and increased in the afternoon with highest values during the night. Calculated values of turgor pressure showed no consistent diurnal trend and did not correlate with the rate of leaf extension. There was no consistent difference in turgor between treatments. Turgor after stress relaxation varied diurnally. The difference between turgors before and after stress relaxation also varied diurnally and was largely in phase with the diurnal pattern of leaf extension. These data are consistent with either a causal role for growth turgor (difference between turgors before and after stress relaxation) in the regulation of cell expansion, or a diurnal variation in turgors after relaxation, attributable to different capacities for cell wall loosening at different times of day. Plastic extensibility of leaf tissue showed no diurnal pattern but consistently higher values were found in high-N than in low-N plants. We conclude that the effects of nitrogen supply on leaf water relations did not limit leaf extension, but that nitrogen supply did affect processes associated with cell wall loosening and enlargement. Nitrogen supply did not affect final values of turgor after relaxation, but it presumably affected the rate at which relaxation proceeded.     

Review of ground-based methods to measure the distribution of biomass in forest canopies

? Context

Ecological research and an effective forest management need accurate information on the structure of the forest canopy to understand the biochemical, physiological and biogeochemical processes within a forest.

? Research question

This paper reviews the currently available instruments for measuring the distribution of biomass within forest canopies. We compare the most well-established approaches and present the different measurable parameters. A special focus lies on the resolution of the obtained data.

? Results

It was found that only 3D laser scanners offer data with the resolution required by ecologists, private landholders, the forest industry and the public to detect trends in tree growth patterns and canopy interactions in all three spatial dimensions. But data validation, data analysis and parameter extraction are still under development, and the price of the instrument is quite high.

? Conclusion

Research should focus on the parameter extraction from terrestrial laser scanner data as this could allow the calculation of functional attributes for different sections of a canopy on a high spatial resolution. It could also help ecologists characterize the structure of forest stands in a quick and precise way.     

Diurnal pattern of leaf extension in Salix viminalis relates to the difference in leaf turgor before and after stress relaxation

Relations between leaf dry weight to leaf area (LWA), leaf nitrogen concentration and irradiance inside a natural canopy were studied in Betula pendula Roth., Corylus avellana L. and Lonicera xylosteum L. In all species, LWA increased with increasing irradiance. Relative variability in LWA was smaller in Betula pendula than in the other two species. In Corylus avellana, LWA also depended on total plant height. Foliar nitrogen concentration (on a dry weight basis) increased with increasing irradiance and LWA in Betula pendula, but decreased in the other two species. The interspecific variation in response to light availability and in nitrogen partitioning may be caused by different light demands or different life forms (trees versus shrubs), or both, of the species examined, and must be considered in contemporary canopy models.     

Horizontal and vertical variations in photosynthetic capacity in a Pinus densiflora crown in relation to leaf nitrogen allocation and acclimation to irradiance

We measured horizontal and vertical gradients of light (rPPFD) along four first-order branches of a Pinus densiflora Sieb. & Zucc. crown, and compared variations in specific leaf area (SLA), needle nitrogen concentration (N), chlorophyll concentration (Chl) and photosynthetic capacity (i.e., maximum rate of carboxylation (V(cmax))) along the two axes. The horizontal gradient of rPPFD along first-order branches was similar in magnitude to the vertical gradient of rPPFD from the upper to the lower crown. None of the measured parameters (i.e., SLA, N, Chl and Vcmax) were strictly proportional to rPPFD, although they were more or less correlated with light when data obtained for all of the crown were pooled (r(2) = 0.31-0.80). The slope of rPPFD against N on an area basis (Narea) for a branch in the middle of the crown orientated northward was significantly greater than the slope for a similar branch orientated southward. Horizontal variations were unrelated to age effects because measurements were all on 1-year-old needles. We conclude that factors other than light (i.e., orientation) may influence N allocation within branches. There was considerably less variation in the relationship of Vcmax to Narea (r2 = 0.58) than in the relationship of Vcmax to rPPFD (r2 = 0.41). Fractional N distribution among components of the photosynthetic machinery was constant within the crown. Together with the relationships between rPPFD and N on a mass basis (r2 = 0.80) and SLA and Vcmax (r2 = 0.60), these findings suggest that most light acclimation in P. densiflora occurs through changes in needle morphology (e.g., SLA) during development.     

Poplar leaf biomass distribution and nitrogen dynamics in a poplar-barley intercropped system in southern Ontario, Canada

The effect of hybrid poplar (Populus spp. clone DN 177) leaf biomass distribution on soil nitrification was investigated in two experiments during the 1993, 1994 and 1995 growing seasons in a poplar-barley (Hordeum vulgare cv. OAC Kippen) intercropping experiment established at Guelph, Ontario, Canada. In experiment 1, poplar was intercropped with barley during all three years and the poplar leaves shed during the fall season were removed from the soil surface during 1993 and 1994. In experiment 2, poplar was intercropped with barley in 1993 and with corn (Zea mays cv. Pioneer 3917) in 1994 an 1995, respectively, and the shed poplar leaves were not removed. In experiment 1, the nitrification rates were lower during 1994 and 1995 when the dropped leaves were removed from the field. The total above-ground biomass of barley within 2.5 m of the tree row was 517, 500 and 450 g×m⁻², respectively during the three years, whereas in the middle of the crop row (4–11 m), the corresponding figures were 491, 484 and 464 g×m–2. Mean nitrification rates, N availability and carbon content were higher in soils close to the poplar tree rows (2.5 m) compared to the corresponding values in the middle of the crop alley (4–11 m from the tree row). In experiment 2, where poplar leaves were not removed from the field, nitrification rates in soils within 2.5 m distance from the poplar row were fairly constant (range 100 to 128 μg 100 g⁻¹ dry soil day⁻¹) during the three years. Results suggest that soil nitrification rates, soil carbon content and plant N uptake adjacent to the poplar tree rows are influenced by poplar leaf biomass input in the preceding year. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of nitrogen fertilization on growth in a Salix viminalis stand using a response surface experimental design

In order to find a nitrogen fertilization regime that is economically feasible in commercial short‐rotation forest stands of basket willow (Salix viminalis L.), a field trial was established on a clay soil near Västerås, central Sweden, in early 1990. A response surface design was used in which three levels of treatment were chosen for each single year: year 1—0, 30, 60, kg N ha^?1; year 2—0, 60, 120, kg N ha^?1; year 3—0, 90, 180, kg N ha^?1; year 4—0, 60, 120, kg N ha^?1. Thirty‐two combinations of these levels, out of 81 possible, were chosen and divided into four blocks. Thus, eight unique treatments were randomized to the plots within each block, providing no replicates at the end of the experimental period. This gave us a more flexible and informative experiment than many of those used earlier with rather few treatments, although they were replicated. After 4 years the willow stand was harvested and the accumulated stem production of each treatment was assessed. The effect of nitrogen fertilization on accumulated stem growth over the experimental period was found to be significant only for nitrogen applied in years 2 and 3, with a somewhat (statistically significant) larger effect in year 2. Growth response to nitrogen fertilization was best approximated with a function including terms with applied amount of nitrogen in years 2 and 3 and, furthermore, a term of interaction with negative coefficient between these years. The interaction between these years was interpreted as the system's ability to recycle nitrogen from one year to another.     

Premature losses of leaf area in response to drought and insect herbivory through a leaf lifespan gradientOA

Implications of the differences in leaf life span are still subject to debate in the field of ecophysiology. Since leaf traits associated with these differences may be decisive for determining the distribution of tree species, this topic is particularly relevant in the context of climate change. This study analyzes the effects of the differences in leaf life span on premature losses of leaf area owing to insect herbivory and to abiotic stress. Loss of leaf area may be an important determinant of...     

Shoot structure and photosynthetic efficiency along the light gradient in a Scots pine canopy

We examined the effects of structural and physiological acclimation on the photosynthetic efficiency of Scots pine (Pinus sylvestris L.) shoots. We estimated daily light interception (DLI) and photosynthesis (DPHOT) of a number of sample shoots situated at different positions in the canopy. Photosynthetic efficiency (epsilon) was defined as the ratio of DPHOT to the potential daily light interception (DLI(ref)) defined as the photosynthetically active radiation (PAR) intercepted per unit area of a sphere at the shoot location. To calculate DLI(ref), DLI and DPHOT, the radiation field surrounding a shoot in the canopy was first modeled using simulated directional distributions of incoming PAR on a clear and an overcast day, and estimates of canopy gap fraction in different directions provided by hemispherical photographs. A model of shoot geometry and measured data on shoot structure and photosynthetic parameters were used to simulate the distribution of PAR irradiance on the needle surface area of the shoot. Photosynthetic efficiency (epsilon) was separated into light-interception efficiency (epsilon(I) = DLI/DLI(ref)) and conversion efficiency (epsilon(PHOT) = DPHOT/DLI). This allowed us to quantify separately the effect of structural acclimation on the efficiency of photosynthetic light capture (epsilon(l)), and the effect of physiological acclimation on conversion efficiency (epsilon(PHOT)). The value of epsilon increased from the top to the bottom of the canopy. The increase was largely explained by structural acclimation (higher epsilon(I)) of the shade shoots. The value of epsilon(PHOT) of shade foliage was similar to that of sun foliage. Given these efficiencies, the clear-day value of DPHOT for a sun shoot transferred to shade was only half that of a shade shoot at its original position. The method presented here provides a tool for quantitatively estimating the role of acclimation in total canopy photosynthesis.     

Application of the pipe model theory to non-destructive estimation of leaf biomass and leaf area of pruned agroforestry trees

The relationship of branch cross sectional area (CS) to leaf biomass (LM) and leaf area (LA) was studied in three agroforestry tree species,Calliandra calothyrsus Maissn.,Erythrina berteroana Urban andErythrina poeppigiana (Walpers) O.F. Cook, to develop a non-destructive method for the estimation of LM and LA for trees managed with periodic pruning. Variation in these relationships was observed according to the bifurcation level and, in theErythrina spp., by clone. All the relationships were linear except the CS-LM relation in small branches ofE. poeppigiana, where it was initially exponential. At main branch level the relationship of CS to LM and LA was linear in all cases but the regression parameter values varied between species and clones, with determination coefficient (R²) 0.88–0.99. It was concluded that the ratio of main branch CS to LM and LA can be used for non-destructive estimation of the latter variables. The method has the additional benefit that the regression parameter value reflects the allocation of dry matter within a tree and, consequently, may give indications about its possible uses in different agroforestry systems.Work carried out at the Centro Agronómico Tropical de Investigación y Enseñanza (CATIE), Turrialba, Costa Rica.     

Variations in bulk leaf carbon isotope discrimination, growth and related leaf traits among three Populus nigra L. populations

The ongoing global change could be an additional threat to the establishment and the long-term survival of Populus nigra L., an emblematic European riparian species. With the general aim of gaining insights into the adaptive potential of this species, we (i) quantified variations within and among three French P. nigra populations for key physiological attributes, i.e., water-use efficiency (assessed from bulk leaf carbon isotope discrimination, Δ(13)C), growth performance and related leaf traits, (ii) examined genotype and population by environment interactions, and (iii) explored the relationship between Δ(13)C and growth. Thirty genotypes were sampled in each of three naturally established populations and grown in two different sites, Orléans (ORL) and Guémené-Penfao (GMN). In ORL, two similar plots were established and different watering regimes were applied in order to test for the drought response. Significant variations were observed for all traits within and among populations irrespective of site and watering. Trait variation was larger within than among populations. The effect of drought was neither genotype- nor population-dependent, contrary to the effect of site. The population ranking was maintained in all sites and watering regimes for the two most complex traits: Δ(13)C and growth. Moreover, these two traits were unrelated, which indicates that (i) water-use efficiency and growth are largely uncoupled in this species, and (ii) the environmental factors driving genetic structuration for Δ(13)C and growth act independently. The large variations found within populations combined with the consistent differences among populations suggest a large adaptive potential for P. nigra.     

Nodule study in Albizia chinensis in relation to nitrogen metabolism,morphology and biomass

Morphology,biomass,nitrate reductase(NR)and nitrogenase activity in Albizia chinensis(Osb.)Merr.nodules were assessed on monthly and seasonal basis for 1 year.Average NR and nitrogenase activity was higher during the rainy season,reaching a maximum in August.Thereafter,activity decreased through autumn and reached a minimum value during winter.Fresh and dry biomass of nodules increased gradually from summer to the rainy season and then started decreasing with the onset of winter as nodules began to senesce.Among four developmental stages of the nodules that correspond to their increasing age,NR and nitrogenase activity remained low in stage 1 nodules and peaked in stage 2.The activity of both enzymes further decreased with increasing age(stage 2 to stage 4).Morphological features such as shape,diameter and color varied considerably among the developmental stages.Stage 1 nodules were cream-colored,oval to heartshaped with smallest average diameter whereas at stage 2,they became bilobed to tetralobed.On the other hand,stage 3 nodules had the largest average diameter and were multilobed in structure.Stage 4 nodules that correspond to the senescing stage were dark brown to black,multilobed,flattened and hollow due to degeneration of nodular tissue.     

Effects of shade on growth, biomass allocation and leaf morphology in European yew (Taxus baccata L.)

The impact of shade on the growth of European yew (Taxus baccata L.) saplings was investigated over a three-year period using artificial shading to simulate four different light regimes (3, 7, 27 and 100 % relative photosynthetic photon flux density, RPPFD). There was no mortality attributable to shading even under the 3 % RPPFD treatment. Increasing shade was positively associated with specific leaf area, leaf length, leaf width and total chlorophyll content, but negatively associated with plant height, stem diameter, total dry weight and root to leaf and shoot ratio. Discoloration of the foliage occurred in plants grown in 100 % RPPFD conditions (resulting in reduced growth rates) and those transferred to 100 % RPPFD conditions after being shade-acclimated for 2 years. Evidence suggests that T. baccata has the ability to regenerate beneath a lighter canopy but beneath denser canopies gap dynamics will play an important role in facilitating successful regeneration and this needs to be reflected in management of natural populations of this declining species.     

Shoot growth patterns in saplings of Cleyera japonica in relation to light and architectural position

To gain further insight into crown development, the influences of shoot architectural position (branch order) and light environment on patterns of shoot growth of Cleyera japonica Thunberg (Theaceae) were investigated. Annual shoot length and light environment were positively correlated within same-order branches. Shoot length differed significantly among branch orders: shoot length was greater for the lower-order branches when light environments were comparable. Lower-order branches lengthened to a certain extent even if light availability was relatively low, whereas higher-order branches did not grow vigorously even when light availability was relatively high. Within same-order branches, branching was independent of the light environment of the shoot. Sylleptic shoot production differed significantly among branch orders, with most sylleptic shoots being produced on second-order branches. It is concluded that both light condition and architectural position of shoots must be considered when examining the mechanisms underlying crown development.     

Spatial distribution of leaf morphological and physiological characteristics in relation to local radiation regime within the canopies of 3-year-old Populus clones in coppice culture

Spatial distributions of leaf characteristics relevant to photosynthesis were compared within high-density coppice canopies of Populus spp. of contrasting genetic origin. We studied three clones representative of the range in growth potential, leaf morphology, coppice and canopy structure: Clone Hoogvorst (Hoo) (Populus trichocarpa Torr. & Gray x Populus deltoides Bartr. & Marsh), Clone Fritzi Pauley (Fri) (Populus trichocarpa Torr. & Gray) and Clone Wolterson (Wol) (Populus nigra L.). Leaf area index ranged from 2.7 (Fri and Wol) to 3.8 (Hoo). The clones exhibited large vertical variation in leaf area density (0.02-1.42 m2 m-3). Leaf dry mass per unit leaf area (DM(A)) increased with increasing light in Clones Hoo and Fri, from about 56 g m-2 at the bottom of the canopy to 162 g m-2 at the top. In Clone Wol, DM(A) varied only from 65 to 100 g m-2, with no consistent relationship with respect to light. Conversely, nitrogen concentration on a mass basis was nearly constant (around 1.3-2.1%) within the canopies of Clones Hoo and Fri, but increased strongly with light in Clone Wol, from 1.4% at the bottom of the canopy to 4.1% at the top. As a result, nitrogen per unit leaf area (N(A)) increased with light in the canopies of all clones, from 0.9 g m-2 at the bottom to 2.9 g m-2 at the top. Although a single linear relationship described the dependence of maximum carboxylation rate (17-93 micromol CO2 m-2 s-1) or electron transport capacity (45-186 micromol electrons m-2 s-1) on N(A), for all clones, Clone Wol differed from Clones Hoo and Fri by exhibiting a higher dark respiration rate at low N(A) (1.8 versus 0.8 micromol CO2 m-2 s-1).     

Soil acidity, and the growth, biomass partitioning and leaf mineral composition of honeylocust (Gleditsia triacanthos L.) seedlings

Honeylocust seedlings (Gleditsia triacanthos L.) were grown in cylinders containing soil adjusted to pH 4, 5 or 6, and harvested every 10 days for 40 days for dry weight and leaf mineral analysis. Total weight of plants grown at pH 4 was less than that of plants grown at pH 5 or 6. Root weight accounted for a greater proportion of total weight in plants grown at pH 4 than in plants grown at pH 5 or 6. Root growth as a function of total plant growth was higher in plants grown at pH 4 than in plants grown at pH 5 or 6, whereas leaf growth as a function of total plant growth was less in plants grown at pH 4 than in plants grown at pH 5 or 6. However, the relationships between root biomass and root length and between leaf biomass and leaf area were the same in all treatments. An analysis of total leaf concentrations of Ca, P, K, Mg, Mn and Al indicated that Al accumulation in leaves was significantly related to a decrease in plant growth at pH 5. A leaf tissue aluminum concentration of 35 microg g(-1) was associated with toxicity symptoms and a 25% reduction in total plant weight.     

Variation in leaf phenolics of field‐cultivated willow (Salix myrsinifolia) clones in relation to occurrence of Melampsora rust

Concentrations of potential antifungal phenolics in field‐cultivated willow (Salix myrsinifolia) clones were analysed during three growing seasons, and correlated to the occurrence of Melampsora rust. Consistent relationships between phenolics and rust were not found across the experimental years. There was significant clonal and temporal variation in phenolic content and rust frequency. Levels of some phenolics varied considerably within a sequence of four full‐grown leaves, but the variation in rust occurrence within the same leaf sequence was nonsignificant. The results suggest that the possible association between willow phenolics and rusts is not straightforward, and emphasize the importance of long‐term studies to investigate the chemical basis of willow rust interactions.     

氮磷配比对水曲柳光合作用的影响

以水曲柳2年生苗木作为试验材料,在人工气候室中用霍格兰水培营养液修改配方(四种氮、磷的不同配比,清水作为对照)进行水培试验,测定了叶片的光合作用、叶绿素荧光以及叶绿素含量等,以期从光合作用角度探讨不同配比的氮、磷对水曲柳幼苗生产力的影响。结果表明:不同氮、磷配比对水曲柳的多个光合指标差异影响显著。在一定范围内随着施磷量的逐渐增加净光合速率(Pn)升高;施磷肥对提高水曲柳叶片光合作用能力有积极作用;磷胁迫使叶绿素含量降低且对水曲柳叶片的碳同化和光能利用产生负影响。     

Leaf nitrogen productivity is the major factor behind the growth reduction induced by long-term salt stress

Plant growth response to salinity on a scale of years has not been studied in terms of growth analysis. To gain insights into this topic, 2-year-old Mediterranean Fan Palm (Chamaerops humilis L.) and Mexican Fan Palm (Washingtonia robusta H. Wendl) seedlings, each with its own distinct plant morphology, were grown for 2 years in a peat soil and irrigated with water of 2 dS m(-1) (control) or 8 dS m(-1) (saline). Plants were harvested on seven occasions and the time trends in relative growth rate (RGR, the rate of increase of biomass per unit of biomass already existing) and its components were analysed. In the long term, salinity produced a slight reduction in the mean RGR, values in both species. In the short term, salinity caused a reduction in RGR. However, during the second year, plants irrigated with 8 dS m(-1) grew somewhat more quickly than the control plants, probably as a result of delay in the growth kinetics due to salinity. Regarding RGR components, leaf nitrogen productivity (the rate of biomass gain per unit leaf N and time) was the major factor causing the differences in RGR resulting from salinity. Washingtonia robusta showed a relatively high plasticity in plant morphology by increasing root and decreasing stem biomass allocation in the presence of salinity. However, the long-term response of W. robusta to salinity, based to a great extent, on this morphological plasticity, was less effective than that of C. humilis, which is based mainly on the contribution of leaf N to RGR values.