Maximizing daily canopy photosynthesis with respect to the leaf nitrogen allocation pattern in the canopy

Summary A model of daily canopy photosynthesis was constructed taking light and leaf nitrogen distribution in the canopy into consideration. It was applied to a canopy of Solidago altissima. Both irradiance and nitrogen concentration per unit leaf area decreased exponentially with increasing cumulative leaf area from the top of the canopy. The photosynthetic capacity of a single leaf was evaluated in relation to irradiance and nitrogen concentration. By integration, daily canopy photosynthesis was calculated for various canopy architectures and nitrogen allocation patterns. The optimal pattern of nitrogen distribution that maximizes the canopy photosynthesis was determined. Actual distribution of leaf nitrogen in the canopy was more uniform than the optimal one, but it realized over 20% more photosynthesis than that under uniform distribution and 4.7% less photosynthesis than that under the optimal distribution. Redeployment of leaf nitrogen to the top of the canopy with ageing should be more effective in increasing total canopy photosynthesis in a stand with a dense canopy than in a stand with an open canopy.     

Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees

Abstract. Seasonal changes in photosynthesis, leaf nitrogen (N) contents and leaf mass per area (LMA) were observed over three growing seasons in open-grown sun-lit leaves of red maple ( Acer rubrum ), sugar maple ( A. sacchamm ) and northern pin oak ( Quereus ellipsoidalis ) trees in southern Wisconsin. Net photosynthesis and leaf N were highly linearly correlated on both mass and area bases within all species from late spring until leaf senescence in fall. Very early in the growing season leaves had high N concentrations, but low photosynthetic rates per unit leaf N, suggesting that leaves were not fully functionally developed at that time. Leaf N per unit area and LMA had nonparallel seasonal patterns, resulting in differing relationships between leaf N/area and LMA in the "early versus late growing season. As a result of differences in seasonal patterns between leaf N/area and LMA, net photosynthesis/area was higher for a given LMA in the spring than fall, and the overall relationships between these two parameters were poor.     

Comparative ecophysiology of leaf and canopy photosynthesis

Leaves and herbaceous leaf canopies photosynthesize efficiently although the distribution of light, the ultimate resource of photosynthesis, is very biased in these systems. As has been suggested in theoretical studies, if a photosynthetic system is organized such that every photosynthetic apparatus photosynthesizes in concert, the system as a whole has the sharpest light response curve and is most adaptive. This condition can be approached by (i) homogenization of the light environment and (ii) acclimation of the photosynthetic properties of leaves or chloroplasts to their local light environments. This review examines these two factors in the herbaceous leaf canopy and in the leaf. Changes in the inclination of leaves in the canopy and differentiation of mesophyll into palisade and spongy tissue contribute to the moderation of the light gradient. Leaf and chloroplast movements in the upper parts of these systems under high irradiances also moderate light gradients. Moreover, acclimation of leaves and chloroplasts to the local light environment is substantial. These factors increase the efficiency of photosynthesis considerably. However, the systems appear to be less efficient than the theoretical optimum. When the systems are optically dense, the light gradients may be too great for leaves or chloroplasts to acclimate. The loss of photosynthetic production attributed to the imperfect adjustment of photosynthetic apparatus to the local light environment is most apparent when the photosynthesis of the system is in the transition between the light-limited and light-saturated phases. Although acclimation of the photosynthetic apparatus and moderation of light gradients are imperfect, these markedly raise the efficiency of photosynthesis. Thus more mechanistic studies on these adaptive attributes are needed. The causes and consequences of imperfect adjustment should also be investigated.     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

Adjustment of leaf photosynthesis to shade in a natural canopy: reallocation of nitrogen

The present study was performed to investigate the adjustment of the constituents of the light and dark reactions of photosynthesis to the natural growth irradiance in the leaves of an overstorey species, Betula pendula Roth, a subcanopy species Tilia cordata P. Mill., and a herb Solidago virgaurea L. growing in a natural plant community in Järvselja, Estonia. Shoots were collected from the site and properties of individual leaves were measured in a laboratory, by applying a routine of kinetic gas exchange and optical measurements that revealed photosystem II (PSII), photosystem I (PSI), and cytochrome b₆f densities per leaf area and the distribution of excitation (or chlorophyll, Chl) between the two photosystems. In parallel, N, Chl and ribulose-bisphosphate carboxylase-oxygenase (Rubisco) content was measured from the same leaves. The amount of N in photosynthetic proteins was calculated from the measured contents of the components of the photosynthetic machinery. Non-photosynthetic N was found as the residual of the budget. Growth in shade resulted in the decrease of leaf dry mass to a half of the DW in sun leaves in each species, but the total variation, from the top to the bottom of the canopy, was larger. Through the whole cross-section of the canopy, leaf dry weight (DW) and Rubisco content per area decreased by a factor of four, N content by a factor of three, but Chl content only by a factor of 1.7. PSII density decreased by a factor of 1.9, but PSI density by a factor of 3.2. The density of PSI adjusted to shade to a greater extent than the density of PSII. In shade, the distribution of N between the components of the photosynthetic machinery was shifted toward light-harvesting proteins at the expense of Rubisco. Non-photosynthetic N decreased the most substantially, from 54% in the sun leaves of B. pendula to 11% in the shade leaves of T. cordata. It is concluded that the redistribution of N toward light-harvesting Chl proteins in shade is not sufficient to keep the excitation rate of a PSII centre invariant. Contrary to PSII, the density of PSI – the photosystem that is in immediate contact with the carbon assimilation system – shade-adjusts almost proportionally with the latter, whereas its Chl antenna correspondingly increases. Even under N deficiency, a likely condition in the natural plant community, a substantial part of N is stored in non-photosynthetic proteins under abundant irradiation, but much less under limiting irradiation. At least in trees the general sequence of down-regulation due to shade adjustment is the following: (1) non-protein cell structures and non-photosynthetic proteins; (2) carbon assimilation proteins; (3) light reaction centre proteins, first PSI; and (4) chlorophyll-binding proteins.     

Relationship between maximum leaf photosynthesis,nitrogen content and specific leaf area in balearic endemic and non-endemic mediterranean species

Gas exchange parameters, leaf nitrogen content and specific leaf area (SLA) were measured in situ on 73 C3 and five C4 plant species in Mallorca, west Mediterranean, to test whether species endemic to the Balearic Islands differed from widespread, non-endemic Mediterranean species and crops in their leaf traits and trait inter-relationships. Endemic species differed significantly from widespread species and crops in several parameters; in particular, photosynthetic capacity, on an area basis (A), was 20 % less in endemics than in non-endemics. Similar differences between endemics and non-endemics were found in parameters such as SLA and leaf nitrogen content per area (Na). Nevertheless, most of the observed differences were found only within the herbaceous deciduous species. These could be due to the fact that most of the non-endemic species within this group have adapted to ruderal areas, while none of the endemics occupies this kind of habitat. All the species-including the crops-showed a positive, highly significant correlation between photosynthetic capacity on a mass basis (Am), leaf nitrogen content on a mass basis (Nm) and SLA. However, endemic species had a lower Am for any given SLA and Nm. Hypotheses are presented to explain these differences, and their possible role in reducing the distribution of many endemic Balearic species is discussed.     

Effect of 3D nitrogen, dry mass per area and local irradiance on canopy photosynthesis within leaves of contrasted heterogeneous maize crops

BACKGROUND AND AIMS: Nitrogen partitioning within stands has been described fairly comprehensively, especially for C(3) plants in dense stands where the horizontal heterogeneity of foliage distribution is relatively small. Nitrogen has been shown to be distributed vertically and in parallel to light, maximizing carbon assimilation and stand productivity. Conversely, row crops such as maize (C(4) plants) are characterized by strong horizontal heterogeneity of foliage distribution, and a three-dimensional (3D) approach is required to investigate the combined effect of spatial distribution of nitrogen and light on canopy photosynthesis. MODEL: The 3D geometry of maize canopies was modelled with varying densities and at different developmental stages using plant digitizing under field conditions. For lamina parts, photosynthesis was measured and nitrogen content per unit area (N(a)) was described from analysis of nitrogen content per unit mass (N(m)) and dry mass per unit area (M(a)). Hyperbolic relationships between photosynthesis at irradiance saturation (P(max)) and N(a) were established as well as a linear relationship between dark respiration (R(d)) and N(a), whereas quantum efficiency (alpha) was found to be independent of N(a). KEY RESULTS AND CONCLUSIONS: N(m), M(a) and N(a) were shown to change over time vertically (i.e. between laminae), which has been largely reported previously, and horizontally (i.e. within laminae), which has scarcely been described previously. Even if M(a) played a major role in N(a), a strong relationship between N(a) and M(a) could not be demonstrated, whereas several previous studies have found that N(a) was essentially related to M(a) rather than N(m). From simulations of radiative exchange using a 3D volume-based approach and lamina photosynthesis using a hyperbola, it was shown that real patterns of N(a) partitioning could increase daily crop photosynthesis by up to 8 % compared with uniform patterns of N(a), especially for the earliest stages of stand development.     

Plastic changes of leaf mass per area and leaf nitrogen content in response to canopy openings in saplings of eight deciduous broad-leaved tree species

Leaf nitrogen content per area (N_area) is a good indicator of assimilative capacity of leaves of deciduous broad-leaved trees. This study examined the degrees of increase in N_area in response to canopy openings as leaf mass per area (LMA) and leaf nitrogen content per mass (N_mass) in saplings of eight deciduous broad-leaved tree species in Hokkaido, northern Japan. Five of the species were well-branched species with a large number of small leaves (lateral-growth type), and the other three species were less-branched species with a small number of large leaves (vertical-growth type). The degrees of increase in N_area were compared between the two crown types. In closed-canopy conditions, leaves of the vertical-growth species tended to have a lower LMA and higher N_mass than those of the lateral-growth species, which resulted in similar N_area for both. LMA increased in canopy openings in the eight species, and the degrees of increase were not largely different between the lateral- and vertical-growth species. On the contrary, N_mass was unchanged in canopy openings in the eight species. As a result, N_area of each species increased in canopy openings in proportion to the increase in LMA, and the degrees of increase in N_area were similar in the lateral- and vertical-growth species. Therefore, this study showed that the degrees of increase in N_area were not correlated with the crown architecture (i.e., the lateral- and vertical-growth types).     

Mean residence time in multicompartmental models with time delays

Calculation of the mean residence time (MRT) of a drug in a stationary compartmental model is classically carried out from several expressions. Nevertheless, one or more time delays between compartments modify the mean residence times. It is the aim of this paper to propose a general method for MRT calculations, in any n-compartmental models which may include time delays. As examples, catenary and mammillary models are considered.     

Monitoring leaf photosynthesis with canopy spectral reflectance in rice

Non-destructive and rapid method for assessment of leaf photosynthetic characteristics is needed to support photosynthesis modelling and growth monitoring in crop plants. We determined the quantitative relationships between leaf photosynthetic characteristics and canopy spectral reflectance under different water supply and nitrogen application rates. The responses of reflectance at red radiation (wavelength 680 nm) to different water contents and nitrogen rates were parallel to those of leaf net photosynthetic rate (P _N). The relationships of reflectance at 680 nm and ratio index of R(810,680) (near infrared/red, NIR/R) to P _N of different leaf positions and leaf layers in rice indicated that the top two full leaves were the best leaf positions for quantitative monitoring of leaf P _N with remote sensing technique, and the ratio index R(810,680) was the best ratio index for evaluating leaf photosynthetic characteristics in rice. Testing of the models with independent data sets indicated that R(810,680) could well estimate P _N of top two leaves and canopy leaf photosynthetic potential in rice, with the root mean square error of 0.25, 0.16, and 4.38, respectively. Hence R(810,680) can be used to monitor leaf photosynthetic characteristics at different growth stages of rice under diverse growing conditions.     

Some quantitative relationships between leaf area index and canopy nitrogen content and distribution

In a previous study (Yin et al. 2000. Annals of Botany 85: 579-585), a generic logarithmic equation for leaf area index (L) in relation to canopy nitrogen content (N) was developed: L=(1/ktn)1n(1+ktnN/nb). The equation has two parameters: the minimum leaf nitrogen required to support photosynthesis (nb), and the leaf nitrogen extinction coefficient (ktn). Relative to nb, there is less information in the literature regarding the variation of ktn. We therefore derived an equation to theoretically estimate the value of ktn. The predicted profile of leaf nitrogen in a canopy using this theoretically estimated value of ktn is slightly more uniform than the profile predicted by the optimum nitrogen distribution that maximizes canopy photosynthesis. Relative to the optimum profile, the predicted profile is somewhat closer to the observed one. Based on the L-N logarithmic equation and the theoretical ktn value, we further quantified early leaf area development of a canopy in relation to nitrogen using simulation analysis. In general, there are two types of relations between L and N, which hold for canopies at different developmental phases. For a fully developed canopy where the lowest leaves are senescing due to nitrogen shortage, the relationship between L and N is described well by the logarithmic model above. For a young, unclosed canopy (i.e. L < 1.0), the relation between L and N is nearly linear. This linearity is virtually the special case of the logarithmic model when applied to a young canopy where its total nitrogen content approaches zero and the amount of nitrogen in its lowest leaves is well above nb. The expected patterns of the L-N relationship are discussed for the phase of transition from young to fully developed canopies.     

The role of nitrogen in a simple scheme to scale up photosynthesis from leaf to canopy

A simple analytical scheme, involving the distribution of nitrogen, to scale up photosynthesis from leaf to canopy is proposed. The scheme is based on the assumption that there are two pools of nitrogen in leaves: nitrogen in photosynthetic, degradable structures (N_p) and nitrogen in non-photosynthetic and non-degradable structures (N_s). The rate of photon-saturated photosynthesis, F_m, is assumed to be proportional to N_p and is distributed inside the canopy similarly to photon flux density (PFD). Prior assumptions of an optimum distribution of nitrogen are not a prerequisite. Calculations made with the scheme lead to development of the hypothesis that the canopy can be treated as a ‘big leaf’ on the time scales involved in acclimation of photosynthesis to PFD. Simulations using parameters for tree species with different requirements for PFD show that shade-tolerant species may have denser canopies than sun-demanding species because of smaller amounts of non-photosynthetic structural nitrogen and/or supporting tissue in their leaves.     

Microgravity effects on thylakoid,single leaf,and whole canopy photosynthesis of dwarf wheat

The concept of using higher plants to maintain a sustainable life support system for humans during long-duration space missions is dependent upon photosynthesis. The effects of extended exposure to microgravity on the development and functioning of photosynthesis at the leaf and stand levels were examined onboard the International Space Station (ISS). The PESTO (Photosynthesis Experiment Systems Testing and Operations) experiment was the first long-term replicated test to obtain direct measurements of canopy photosynthesis from space under well-controlled conditions. The PESTO experiment consisted of a series of 21–24 day growth cycles of Triticum aestivum L. cv. USU Apogee onboard ISS. Single leaf measurements showed no differences in photosynthetic activity at the moderate (up to 600 μmol m⁻² s⁻¹) light levels, but reductions in whole chain electron transport, PSII, and PSI activities were measured under saturating light (>2,000 μmol m⁻² s⁻¹) and CO₂ (4000 μmol mol⁻¹) conditions in the microgravity-grown plants. Canopy level photosynthetic rates of plants developing in microgravity at ∼280 μmol m⁻² s⁻¹ were not different from ground controls. The wheat canopy had apparently adapted to the microgravity environment since the CO₂ compensation (121 vs. 118 μmol mol⁻¹) and PPF compensation (85 vs. 81 μmol m⁻² s⁻¹) of the flight and ground treatments were similar. The reduction in whole chain electron transport (13%), PSII (13%), and PSI (16%) activities observed under saturating light conditions suggests that microgravity-induced responses at the canopy level may occur at higher PPF intensity.     

Physiological comparisons of two soybean cultivars differing in canopy photosynthesis. II. Variation in specific leaf weight,nitrogen, and protein components

Cultivar differences in canopy apparent photosynthesis (CAP) have been observed in soybean (Glycine max (L.) Merr.) but little is known about the physiological mechanisms which are responsible for such differences. This study was initiated to determine if variation in ribulose 1,5-bisphosphate carboxylase (RuBPCase) and soluble protein exists among cultivars which differ in CAP during reproductive growth. In addition, the relationship between specific leaf weight (SLW) and leaf protein was examined. Two Maturity Group VI cultivars, Tracy (high CAP) and Davis (low CAP), were grown in the field during 1979, 1980, and 1981 and in a greenhouse experiment. Leaves located at two canopy positions (topmost, fully expanded leaf and eighth node from the top) in 1979 and three canopy positions (those mentioned, plus the fourth node from the top) in 1980 and 1981 were sampled. Leaves at the two upper canopy positions exhibited greater SLW, RuBPCase m^–2, and soluble protein m^–2 than found at the eighth node down. Photosynthetic capacity of leaves at inner canopy regions was therefore affected by both light penetration into the canopy and leaf protein status. Over the three year period, the SLW was 23 percent and the soluble protein m^–2 leaf 21 percent greater in Tracy than in Davis. Although the trend in RuBPCase m^–2 leaf was not significant, it was consistently greater in Tracy in the field and greenhouse. No cultivar differences were observed when the proteins were expressed on a unit of leaf dry weight. The quantity of RuBPCase per unit leaf area was positively correlated with SLW with significant partial correlation coefficients of 0.62, 0.67, 0.35, and 0.82 for 1979, 1980, 1981, and the greenhouse study, respectively. Since these cultivars have similar leaf area indices during September, the greater SLW of Tracy is translated into more photosynthetic proteins per unit ground area and higher CAP rate.Abbreviations AP Leaf Apparent Photosynthesis - CAP Canopy Apparent Photosynthesis - DAP Days After Planting - DTT Dithothreitol - HEPES N-2-hydroethylpiperazine N-2 ethanesulfonic acid - LAI Leaf Area Index - LSD Least Significant Difference - PPFD Photosynthetic Photon Flux Density - PVP-40 Polyvinylpolypyrroledone (molecular weight, 4000) - RuBPCase Ribulose 1,5-bisphosphate Carboxylase - SLW Specific Leaf Weight     

Mean labor time of a leaf

The mean labor time of a leaf (hour/day^–1) is defined as the ratio of mean daily photosynthetic rate of a leaf (D_a; molm^–2day^–1) to the mean value of potential hourly photosynthetic rate (6060A_max mol m^–2h^–1) of the leaf. A model was proposed to estimate mean labor time of leaves. Mean labor time was obtained as the product of 24 (hours/day^–1) and the four effects, each of which reduces production of a leaf: diel change in light (Diel Effect), reduction in light during cloudy and rainy days (Cloudy Effect), shading on the focal leaves (Shading Effect), and midday and afternoon depression in photosynthesis (Depression Effect). These four effects were estimated for open grown saplings of alder (Alnus sieboldiana), by measuring instantaneous photosynthetic rate and photon flux density above each leaf. The potential daily photosynthetic rate calculated from diel light condition in a clear day was 46.5% of hypothetical daily photosynthetic rate where maximum instantaneous photosynthetic rate was assumed to last throughout the life of the leaf (Diel Effect). The average of the daily photosynthetic rate considering clear, cloudy and rainy days was 79.7% of the clear day (Cloudy Effect). The photosynthetic rate estimated from light condition on the leaf was 85.6% of that in the open site (Shading Effect). Midday depression reduced the daily photosynthetic rate to 72.1% of the potential daily photosynthetic rate (Depression Effect). The product of the four effects multiplied by 24h gave the estimate of mean labor time of leaves to be approximately 5.5 (h/day^–1).     

Coordination theory of leaf nitrogen distribution in a canopy

It has long been observed that leaf nitrogen concentrations decline with depth in closed canopies in a number of plant communities. This phenomenon is generally believed to be related to a changing radiation environment and it has been suggested by some researchers that plants allocate nitrogen in order to optimize total whole canopy photosynthesis. Although optimization theory has been successfully utilized to describe a variety of physiological and ecological phenomena, it has some shortcomings that are subject to criticism (e.g., time constraints, oversimplifications, lack of insights, etc.). In this paper we present an alternative to the optimization theory of plant canopy nitrogen distribution, which we term coordination theory. We hypothesize that plants allocate nitrogen to maintain a balance between two processes, each of which is dependent on leaf nitrogen content and each of which potentially limits photosynthesis. These two processes are defined as W_c, the Rubiscolimited rate of carboxylation, and W_j, the electron transport-limited rate of carboxylation. We suggest that plants allocate nitrogen differentially to, leaves in different canopy layers in such a way that W_c and W_j remain roughly balanced. In this scheme, the driving force for the allocation of nitrogen within a canopy is the difference between the leaf nitrogen content that is required to bring W_c and W_j into balance and the current nitrogen content. We show that the daily carbon assimilation of a canopy with a nitrogen distribution resulting from this internal coordination of W_c and W_j is very similar to that obtained using optimization theory.     

Effects of seasonal and interannual variations in leaf photosynthesis and canopy leaf area index on gross primary production of a cool-temperate deciduous broadleaf forest in Takayama, Japan

Revealing the seasonal and interannual variations in forest canopy photosynthesis is a critical issue in understanding the ecological mechanisms underlying the dynamics of carbon dioxide exchange between the atmosphere and deciduous forests. This study examined the effects of temporal variations of canopy leaf area index (LAI) and leaf photosynthetic capacity [the maximum velocity of carboxylation (V _cmax)] on gross primary production (GPP) of a cool-temperate deciduous broadleaf forest for 5 years in Takayama AsiaFlux site, central Japan. We made two estimations to examine the effects of canopy properties on GPP; one is to incorporate the in situ observation of V _cmax and LAI throughout the growing season, and another considers seasonality of LAI but constantly high V _cmax. The simulations indicated that variation in V _cmax and LAI, especially in the leaf expansion period, had remarkable effects on GPP, and if V _cmax was assumed constant GPP will be overestimated by 15%. Monthly examination of air temperature, radiation, LAI and GPP suggested that spring temperature could affect canopy phenology, and also that GPP in summer was determined mainly by incoming radiation. However, the consequences among these factors responsible for interannual changes of GPP are not straightforward since leaf expansion and senescence patterns and summer meteorological conditions influence GPP independently. This simulation based on in situ ecophysiological research suggests the importance of intensive consideration and understanding of the phenology of leaf photosynthetic capacity and LAI to analyze and predict carbon fixation in forest ecosystems.     

Specific leaf area relates to the differences in leaf construction cost,photosynthesis, nitrogen allocation,and use efficiencies between invasive and noninvasive alien congeners

Comparisons between invasive and native species may not characterize the traits of invasive species, as native species might be invasive elsewhere if they were introduced. In this study, invasive Oxalis corymbosa and Peperomia pellucida were compared with their respective noninvasive alien congeners. We hypothesized that the invasive species have higher specific leaf (SLA) than their respective noninvasive alien congeners, and analyzed the physiological and ecological consequences of the higher SLA. Higher SLA was indeed the most important trait for the two invaders, which was associated with their lower leaf construction cost, higher nitrogen (N) allocation to photosynthesis and photosynthetic N use efficiency (PNUE). The higher N allocation to photosynthesis of the invaders in turn increased their PNUE, N content in photosynthesis, biochemical capacity for photosynthesis, and therefore light-saturated photosynthetic rate. The above resource capture-, use- and growth-related traits may facilitate the two invaders' invasion, while further comparative studies on a wider range of invasive and noninvasive congeners are needed to understand the generality of this pattern and to fully assess the competitive advantages afforded by these traits.     

Anatomical basis of the change in leaf mass per area and nitrogen investment with relative irradiance within the canopy of eight temperate tree species

Changes in leaf mass per area (LMA), nitrogen content on a mass-basis (N_m) and on an area basis (N_a) with relative irradiance were assessed in leaves of eight temperate species harvested at different depths in a canopy. Relative irradiance (GSF) at the points of leaf sampling was estimated by hemispheric photographs. There was a strong species-dependent positive relationship between LMA and GSF for all species. Shade-tolerant species such as Fagus sylvatica showed lower LMA for the same GSF than less tolerant species as Quercus pyrenaica or Quercus petraea. The only evergreen species in the study, Ilex aquifollium, had the highest LMA, independent of light environment, with minimum values much higher than the rest of the broad-leaved species studied. There was no relation between N_m and GSF for most species studied and only a very weak relation for the relative shade-intolerant species Q. pyrenaica. Within each species, the pattern of N_a investment with regard to GSF was linked mainly to LMA. At the same relative irradiance, differences in N_a among species were conditioned both by the LMA–GSF relationship and by the species N_m value. The lowest N_m value was measured in I. aquifollium (14.3 ± 0.6 mg g^–1); intermediate values in Crataegus monogyna (16.9 ± 0.6 mg g^–1) and Prunus avium (19.1 ± 0.6 mg g^–1) and higher values, all in a narrow range (21.3 ± 0.6 to 23 ± 0.6 mg g^–1), were measured for the other five species. Changes in LMA with the relative irradiance were linked both to lamina thickness (LT) and to palisade/spongy parenchyma ratio (PP/SP). In the second case, the LMA changes may be related to an increase in lamina density as palisade parenchyma involves higher cell packing than spongy parenchyma. However, since PP/SP ratio showed a weak species-specific relationship with LMA, the increase in LT should be the main cause of LMA variation.