Systematics of the Late Carboniferous Medullosalean Pteridosperm Laveineopteris and Its Associated Cyclopteris Leaves

Laveineopteris is emended to become a morphogenus for the vegetative parts of certain medullosalean pteridosperms that grew in the Westphalian (Late Carboniferous) tropical rainforests. The plant bore pinnate foliage as sun leaves and entire orbicular leaves (cyclopterids) as shade leaves. The cyclopterids were attached to the axis of the saplings and epiphyllously to the lower parts of the pinnate fronds. Species recognition is mainly by the macro–morphology of the pinnate fronds. However, species can also be distinguished based on the epidermal anatomy of both pinnate fronds and cyclopterids, in particular the distribution and detailed structure of the stomata and hydathodes. Laveineopteris first appeared in the lowland swamp forests of Europe during the early Langsettian (Westphalian A), and there were two principal species: L. loshii and L. tenuifolia . During the early Bolsovian (early Westphalian C), a variety of L. tenuifolia appeared that was adapted to both the lowland swamps, and the upland swamps of the intramontane basins (central Bohemia and Saar–Lorraine). The smaller–pinnuled L. rarinervis also first appeared in the Bolsovian and seems to have been restricted to the lowland swamps. The morphogenus Cyclopteris should be used only for cyclopterid fossils, where they cannot be related to the species of Laveineopteris , Callipteridium or Margaritopteris that originally bore them.     

Colonization Strategies of Two Liana Species in a Tropical Dry Forest Canopy

Lianas impose intense resource competition for light in the upper forest canopy by displaying dense foliage on top of tree crowns. Using repeated access with a construction crane, we studied the patterns of canopy colonization of the lianas Combretum fruticosum and Bonamia trichantha in a Neotropical dry forest in Panama. Combretum fruticosum flushed leaves just before the rainy season, and its standing leaf area quickly reached a peak in the early rainy season (May–June). In contrast, B. trichantha built up foliage area continuously throughout the rainy season and reached a peak in the late rainy season (November). Both species displayed the majority of leaves in full sun on the canopy surface, but C. fruticosum displayed a greater proportion of leaves (26%) in more shaded microsites than B. trichantha (12%). Self-shading within patches of liana leaves within the uppermost 40–50 cm of the canopy reduced light levels measured with photodiodes placed directly on leaves to 4–9 percent of light levels received by sun leaves. Many leaves of C. fruticosum acclimated to shade within a month following the strongly synchronized leaf flushing and persisted in deep shade. In contrast, B. trichantha produced short-lived leaves opportunistically in the sunniest locations. Species differences in degree of shade acclimation were also evident in terms of structural (leaf mass per area, and leaf toughness) and physiological characters (nitrogen content, leaf life span, and light compensation point). Contrasting leaf phenologies reflect differences in light exploitation and canopy colonization strategies of these two liana species.     

Phenotypic plasticity in response to light in the coffee tree

Phenotypic plasticity to light availability was examined at the leaf level in field-grown coffee trees (Coffea arabica). This species has been traditionally considered as shade-demanding, although it performs well without shade and even out-yields shaded coffee. Specifically, we focused our attention on the morpho-anatomical plasticity, the balance between light capture and excess light energy dissipation, as well as on physiological traits associated with carbon gain. A wide natural light gradient, i.e., a diurnal intercepted photon irradiance differing by a factor of 25 between the deepest shade leaves and the more exposed leaves in the canopy, was explored. Responses of most traits to light were non-linear, revealing the classic leaf sun vs. leaf shade dichotomy (e.g., compared with sun leaves, shade leaves had a lower stomatal density, a thinner palisade mesophyll, a higher specific leaf area, an improved light capture, a lower respiration rate, a lower light compensating point and a limited capacity for photoprotection). The light-saturated rates of net photosynthesis were higher in sunlit than in shade leaves, although sun leaves were not efficient enough to use the extra light supply. However, sun leaves showed well-developed photoprotection mechanisms in comparison to shade leaves, which proved sufficient for avoiding photoinhibition. Specifically, a higher non-photochemical quenching coefficient was found in parallel to increases in: (i) zeaxanthin pools, (ii) de-epoxidation state of the xanthophyll cycle, and (iii) activities of some antioxidant enzymes. Intracanopy plasticity depended on the suite of traits considered, and was high for some physiological traits associated with photoprotection and maintenance of a positive carbon balance under low light, but low for most morpho-anatomical features. Our data largely explain the successful cultivation of the coffee tree in both exposed and shade environments, although with a poor resource-use efficiency in high light.     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance

Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M _A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F _L) modify light harvesting significantly. F _L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F _L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M _A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M _A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M _A, reductions in F _L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.     

Scaling carbon dioxide and water vapour exchange from leaf to canopy in a deciduous forest. I. Leaf model parametrization

In order to parametrize a leaf submodel of a canopy level gas-exchange model, a series of photosynthesis and stomatal conductance measurements were made on leaves of white oak (Quercus alba L.) and red maple (Acer rubrum L.) in a mature deciduous forest near Oak Ridge, TN. Gas-exchange characteristics of sun leaves growing at the top of a 30 m canopy and of shade leaves growing at a depth of 3–4 m from the top of the canopy were determined. Measured rates of net photosynthesis at a leaf temperature of 30°C and saturating photosynthetic photon flux density, expressed on a leaf area basis, were significantly lower (P = 0.01; n = 8) in shade leaves (7.9μmol m^?2 s^?1) than in sun leaves (11–5μmol m^?2 s^?1). Specific leaf area increased significantly with depth in the canopy, and when photosynthesis rates were expressed on a dry mass basis, they were not significantly different for shade and sun leaves. The percentage leaf nitrogen did not vary significantly with height in the canopy; thus, rates expressed on a per unit nitrogen basis were also not significantly different in shade and sun leaves. A widely used model integrating photosynthesis and stomatal conductance was parametrized independently for sun and shade leaves, enabling us to model successfully diurnal variations in photosynthesis and evapotranspiration of both classes of leaves. Key photosynthesis model parameters were found to scale with leaf nitrogen levels. The leaf model parametrizations were then incorporated into a canopy-scale gas-exchange model that is discussed and tested in a companion paper (Baldocchi & Harley 1995, Plant, Cell and Environment 18, 1157–1173).     

Effects of the light environment on oviposition preference and survival of a leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae), on Quercus alba L.

Abstract.  1. Oviposition preference and the survival and mortality of larvae of Cameraria hamadryadella (Lepidoptera) on the foliage of Quercus alba L. (Fagaceae) in sun and shade were examined. To separate the effects of the environment of foliation from that of rearing, C. hamadryadella were allowed to oviposit on oak saplings that foliated either in the sun or shade. Subsequently, trees were placed into the sun or shade and larval survival was monitored.
2. Trees that foliated in the shade had leaves that were 30% longer than those that foliated in the sun. However, the density of leaf mines did not differ between trees that foliated in the sun or shade.
3. Larval survival was higher in the shade than in the sun, but no difference in survival due to the site of foliation was detected. Differences in larval survival between the sun and shade environments could not be attributed to differences in natural enemy mortality. However, 'death by other causes' was higher for larvae reared in the sun than in the shade. In sunny environments photochemical processes may reduce foliage quality, which results in lower survival of C. hamadryadella larvae.
4. Ovipositing C. hamadryadella do not appear to discriminate between foliage produced in the sun or shade. This lack of discrimination, coupled with the fact that ovipositing C. hamadryadella are active after dark, accounts for the absence of a difference in abundance between sun and shade.
5. The results presented here, combined with other published studies, argue that different patterns of insect herbivory and abundance in relation to the light level experienced by host-plant foliage might be expected for species that oviposit either in the night or in the day.     

The nocturnal larvae of a specialist folivore perform better on Chromolaena odorata leaves from a shaded environment

Increasing evidence suggests that individuals of the same plant species occurring in different microhabitats often show a degree of phenotypic and phytochemical variation. Consequently, insect herbivores associated with such plant species must deal with environment‐mediated changes or variability in the traits of their host plants. In this study, we examined the effects of habitat condition (shaded vs. full‐sun habitats) on plant traits and leaf characteristics of the invasive alien plant, Chromolaena odorata (L.) King & Robinson (Asteraceae). In addition, the performance was evaluated in two generations of a specialist folivore, Pareuchaetes insulata (Walker) (Lepidoptera: Erebidae: Arctiinae), on leaves obtained from both shaded and full‐sun habitats. The study was done in an area where the insect was introduced as a biological control agent. Leaves growing in shade were less tough, had higher water and nitrogen content, and lower total non‐structural carbohydrate, compared with leaves growing in full sun. Plants growing in shade had longer leaves and were taller, but above‐ground biomass was significantly reduced compared with plants growing in full sun. In both generations (parents and offspring), P. insulata developed faster and had larger pupal mass, increased growth rate, and higher fecundity when reared on shaded foliage compared with full‐sun foliage. Although immature survival and adult longevity did not differ between habitats, Maw's host suitability index indicated that shaded leaves were more suitable for the growth and reproduction of P. insulata. We suggest that the benefits obtained by P. insulata feeding on shaded foliage are associated with reduced toughness and enhanced nitrogen and water content of leaves. These results demonstrate that light‐mediated changes in plant traits and leaf characteristics can affect insect folivore performance.     

Pigment apparatus in <Emphasis Type="Italic">Ajuga reptans</Emphasis> plants as affected by adaptation to light growth conditions

Mechanisms of adaptation of the photosynthetic apparatus at the level of pigment complex in a shade-tolerant bugle plant (Ajuga reptans L.) grown at full solar irradiation in an open plot were studied. In “sun” plants, the content of photosynthetic pigments decreased markedly as compared to “shade” plants grown under a forest canopy at 5–10% of the full solar irradiation. In leaves of sun plants, the portion of β-carotene and lutein in the carotenoid spectrum was higher than in shade plant leaves, antheraxanthin and zeaxanthin were present, and de-epoxidation of violaxanthin was by an order of magnitude higher in sun plant leaves reaching 40%. The data obtained indicate the role of the violaxanthin cycle in the protection of photosynthetic apparatus in a shade-tolerant plant against destruction under excessive irradiation.     

Beech leaf colonization by the endophyte Apiognomonia errabunda dramatically depends on light exposure and climatic conditions

Ozone and light effects on endophytic colonization by Apiognomonia errabunda of adult beech trees (Fagus sylvatica) and their putative mediation by internal defence compounds were studied at the Kranzberg Forest free-air ozone fumigation site. A. errabunda colonization was quantified by "real-time PCR" (QPCR). A. errabunda-specific primers allowed detection without interference by DNA from European beech and several species of common genera of plant pathogenic fungi, such as Mycosphaerella, Alternaria, Botrytis, and Fusarium. Colonization levels of sun and shade leaves of European beech trees exposed either to ambient or twice ambient ozone regimes were determined. Colonization was significantly higher in shade compared to sun leaves. Ozone exhibited a marginally inhibitory effect on fungal colonization only in young leaves in 2002. The hot and dry summer of 2003 reduced fungal colonization dramatically, being more pronounced than ozone treatment or sun exposure. Levels of soluble and cell wall-bound phenolic compounds were approximately twice as high in sun than in shade leaves. Acylated flavonol 3- O-glycosides with putatively high UV-B shielding effect were very low in shade canopy leaves. Ozone had only a minor influence on secondary metabolites in sun leaves. It slightly increased kaempferol 3- O-glucoside levels exclusively in shade leaves. The frequently prominent hydroxycinnamic acid derivative, chlorogenic acid, was tested for its growth inhibiting activity against Apiognomonia and showed an IC50 of approximately 8 mM. Appearance of Apiognomonia-related necroses strongly correlated with the occurrence of the stress metabolite, 3,3',4,4'-tetramethoxybiphenyl. Infection success of Apiognomonia was highly dependent on light exposure, presumably affected by the endogenous levels of constitutive phenolic compounds. Ozone exerted only minor modulating effects, whereas climatic factors, such as pronounced heat periods and drought, were dramatically overriding.     

不同光环境下桂花幼苗的构型差异

通过对不同光照条件下桂花幼苗的冠形、分枝率、叶片在树冠中的空间分布等特征进行研究,结果表明桂花幼苗构型发生了明显的可塑性适应:其树冠对光照条件的变化有显著的可塑性响应。在林隙中的幼苗受光的间歇性影响,总体分枝率明显小于全光、林冠下的幼苗分枝率。全光的幼苗叶片集中于二级枝,叶片长度和叶片面积相对较小,对光照利用充分;而林隙中的幼苗叶片集中于一级枝,避免处于植冠内侧受到遮蔽,表现出较大的叶片长度和叶面积;林冠下的叶片较均匀分布在一、二级枝上,叶片总数量较少,枝条高生长较全光下明显。幼苗在总体分枝格局中表现出独自的特点,即强光环境下产生短枝和高分枝率,在适度庇荫条件下产生长枝及低分枝率,在强度庇荫条件下以较长枝和较高分枝率来同时满足高生长和横向生长的需求。     

Instantaneous canopy photosynthesis: analytical expressions for sun and shade leaves based on exponential light decay down the canopy and an acclimated non-rectangular hyperbola for leaf photosynthesis

Analytical expressions for the contributions of sun and shade leaves to instantaneous canopy photosynthesis are derived. The analysis is based on four assumptions. First, that the canopy is closed in the sense that it is horizontally uniform. Secondly, that there is an exponential profile of light down the canopy with the same decay constant for light from different parts of the sky. Thirdly, that the leaf photosynthetic response to incident irradiance can be described by a three-parameter non-rectangular hyperbola (NRH). And lastly, that light acclimation at the leaf level occurs in only one parameter of the NRH, that describing the light-saturated photosynthetic rate, which is assumed to be proportional to the local averaged leaf irradiance. These assumptions have been extensively researched empirically and theoretically and their limitations are quite well understood. They have been widely used when appropriate. Combining these four assumptions permits the derivation of algebraic expressions for instantaneous canopy photosynthesis which are computationally efficient because they avoid the necessity for numerical integration down the canopy. These are valuable for modelling plant and crop ecosystems, for which canopy photosynthesis is the primary driver. Ignoring the sun/shade dichotomy can result in overestimates of canopy photosynthesis of up to 20 %, but using a rectangular hyperbola instead of a non-rectangular hyperbola to estimate canopy photosynthesis taking account of sun and shade leaves can lead to a similarly sized underestimate.     

Effects of canopy shade on the lipid composition of soybean leaves

The effect of canopy shade on leaf lipid composition was examined in soybeans ( Glycine max cv. Young) grown under field conditions. Expanding leaves were tagged at 50, 58 and 65 days after planting (DAP) in plots with either a high (10 plants m⁻¹ row) or low (1 plant m⁻¹ row) plant density. At 92 DAP, light conditions ranged from a pho-tosynthetic photon flux density (PPFD) of 87% of full sun with a far-red/red (735 nm/645 nm) ratio of 0.9 at upper canopy leaves to extreme shade where the PPFD was 10% of full sun with a far-red/red ratio greater than 6. Highly shaded leaves in the high plant density treatment accumulated triacylglycerol (TG) up to 25% of total leaf lipid, a 2.4-fold increase in TG on a chlorophyll basis compared to leaves in the upper canopy. Although total polar lipid content was reduced up to 50% in shaded leaves, shade had little affect on the lipid content or composition of thylakoid membranes. Shade did not affect leaf chlorophyll content. Therefore, the changes in leaf lipid composition were not related to senescence. These findings suggest that conditions of low irradiance and/or a high FR/R ratio cause a shift in carbon metabolism toward the accumulation of TG, a storage lipid. Eighteen-carbon fatty acid desaturation was also affected in highly shaded leaves where a reduction in linolenic acid (18:3) content was accompanied by a proportional increase in oleic (18:1) and linoleic (18:2) acids.     

The effects of light acclimation during and after foliage expansion on photosynthesis ofAbies amabilis foliage within the canopy

Variation in the photosynthetic function ofAbies amabilis foliage within a canopy was examined and related to three different processes that affect foliage function: foliage aging, sun-shade acclimation that occurred while foliage was expanding, and reacclimation after expansion was complete. Foliage produced in the sun had higher photosynthesis at light saturation (A _max, mol·m^-2·s^-1), dark respiration (mol·m^-2·s^-1), nitrogen content (g·m^-2), chlorophyll content (g·m^-2), and chlorophylla:b ratio, and a lower chlorophyll to nitrogen ratio (chl:N), than foliage produced in the shade. As sun foliage becomes shaded, it becomes physiologically similar to shade foliage, even though it still retains a sun morphology. Shaded sun foliage exhibited lowerA _max, dark respiration, nitrogen content, and chlorophylla:b ratio, and a higher chl:N ratio than sun foliage of the same age remaining in the open. However, shaded sun foliage had a higher chlorophyll content than sun foliage remaining in the open, even though true shade foliage had a lower chlorophyll content than sun foliage. This anomaly arises because as sun foliage becomes shaded, it retains a higher nitrogen content than shade foliage in a similar light environment, but the two forms have similar chl:N ratios. Within the canopy, most physiological indicators were more strongly correlated with the current light environment than with foliage age or leaf thickness, with the exception of chlorophyll content.A _max decreased significantly with both decreasing current light environment of the foliage and increasing foliage age. The same trend with current light and age was found for the chlorophylla:b ratio. Foliage nitrogen content also decreased with a decrease in current light environment, but no distinct pattern was found with foliage age. Leaf thickness was also important for predicting leaf nitrogen content: thicker leaves had more nitrogen than thinner leaves regardless of light environment or age. The chl:N ratio had a strong negative correlation with the current light environment, and, as with nitrogen content, no distinct pattern was found with foliage age. Chlorophyll content of the foliage was not well correlated with any of the three predictor variables: current light environment, foliage age or leaf thickness. On the other hand, chlorophyll content was positively correlated with the amount of nitrogen in a leaf, and once nitrogen was considered, the current light environment was also highly significant in explaining the variation in chlorophyll content. It has been suggested that the redistribution of nitrogen both within and between leaves is a mechanism for photosynthetic acclimation to the current light environment. Within theseA. amabilis canopies, both leaf nitrogen and the chl:N ratio were strongly correlated with the current light environment, but only weakly with leaf age, supporting the idea that changing light is the driving force for the redistribution of nitrogen both within and between leaves. Thus, our results support previous theories on nitrogen distribution and partitioning. However,A _max was significantly affected by both foliage age and the current light environment, indicating that changes in light alone are not enough to explain changes inA _max with time.     

Irradiance heterogeneity within crown affects photosynthetic capacity and nitrogen distribution of leaves in Cedrela sinensis

Because light conditions in the forest understory are highly heterogeneous, photosynthetic acclimation to spatially variable irradiance within a crown is important for crown‐level carbon assimilation. The effect of variation in irradiance within the crown on leaf nitrogen content and photosynthetic rate was examined for pinnate compound leaves in saplings of Cedrela sinensis, a pioneer deciduous tree. Five shading treatments, in which 0, 25, 50, 75 and 100% of leaves were shaded, were established by artificial heavy shading using shade screen umbrellas with 25% transmittance. Although the nitrogen content of leaves was constant regardless of shading treatment, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) content and light‐saturated photosynthetic capacity were lower in shade leaves within partially shaded crowns than within fully shaded crowns. Shade leaves within partially shaded crowns contained higher amount of amino acids. Most shade leaves died in partially shaded crowns, whereas more than half of shade leaves survived in totally shaded crowns. Assumptions on photosynthetic acclimation to local light conditions cannot explain why shade leaves have different photosynthetic capacities and survival rates in between partially and totally shaded crowns. Irradiance heterogeneity within the crown causes a distinct variation in photosynthetic activity between sun and shaded leaves within the crown.     

Coupled response of stomatal and mesophyll conductance to light enhances photosynthesis of shade leaves under sunflecks

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO₂ diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A_n), stomatal conductance (g_s) and mesophyll conductance (g_m) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower A_n, g_m, leaf nitrogen and photosynthetic capacity (A_max) but g_s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g_s and g_m increased, and A_n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of g_m with increased light enables shade leaves to respond quickly to sunflecks.     

Morphological plasticity,photosynthesis and chlorophyll fluorescence of <Emphasis Type="Italic">Athyrium pachyphlebium</Emphasis> at different shade levels

Athyrium pachyphlebium C. is a popular ornamental fern with considerable shade tolerance. The aim of this study was to investigate how the mature sporophytes acclimate to different light levels and to obtain an optimal light environment for their growth both in natural forest canopy and in urban landscapes. Plant growth and morphology, photosynthetic light-response curves and chlorophyll (Chl) fluorescence were measured at four different light levels (45% full sunlight, 30%, 20% and 8%). As the light intensities declined from 45% to 20%, seedling height, crown growth, foliage number and plant lifespan increased significantly. Seedlings grown at 20% light level were vigorous with great ornamental value. Plants grown in deep shade (8% light) showed severe symptoms of lodging and in 45% full sun, the plants showed highlight-stress symptoms. Seedlings in high light levels exhibited a higher light-saturated photosynthetic rate (P _max), light compensation point (LCP), light saturation point (LSP) and a reduced ability for nonphotochemical quenching (NPQ) of excess light than those in low light levels. However, seedlings in low light exhibited greater efficiency in absorbing and utilizing light energy, characterized by higher chlorophyll b (Chl b) and electron transport rate (ETR). These results indicated that a light level of about 20% full sun appeared to be optimal for A. pachyphlebium when both physiological and morphological performance in the landscape were considered.     

Photosynthetic responses of sun- and shade-grown barley leaves to high light: is the lower PSII connectivity in shade leaves associated with protection against excess of light?

In this study, we have compared photosynthetic performance of barley leaves (Hordeum vulgare L.) grown under sun and shade light regimes during their entire growth period, under field conditions. Analyses were based on measurements of both slow and fast chlorophyll (Chl) a fluorescence kinetics, gas exchange, pigment composition; and of light incident on leaves during their growth. Both the shade and the sun barley leaves had similar Chl a/b and Chl/carotenoid ratios. The fluorescence induction analyses uncovered major functional differences between the sun and the shade leaves: lower connectivity among Photosystem II (PSII), decreased number of electron carriers, and limitations in electron transport between PSII and PSI in the shade leaves; but only low differences in the size of PSII antenna. We discuss the possible protective role of low connectivity between PSII units in shade leaves in keeping the excitation pressure at a lower, physiologically more acceptable level under high light conditions.     

The induction of sun and shade leaves of the European beech (Fagus sylvatica L.): anatomical studies

Summary Primordia from buds of sun and shade twigs of European beech (Fagus sylvatica L.) were collected six times a year for anatomical investigations. Differentiation into sun-leaf and shade-leaf primordia was first observed in early August. Sun-leaf primordia had five, and shade-leaf primordia four layers of mesophyll meristem cells. With potted graft unions of beeches possible structural changes of leaf primordia were investigated. Trees adapted to shade develop sun-leaf primordia when put into full daylight, provided the transfer happened before July. Trees adapted to full daylight developed leaf primordia which remained structurally sun-leaf primordia when the plant was kept under shade conditions. Shadeleaf branches of young beech trees cut in February in order to expose the shade buds to full daylight developed either shade leaves or intermediate shade/sun leaves. These experiments show that the subtending leaf may provide the developing axillary bud with photoassimilates, but its character, whether sun or shade leaf, has no influence on the character of the developing leaf primordia.     

Effects of light availability versus hydraulic constraints on stomatal responses within a crown of silver birch

Responses of leaf conductance (g_L) to variation in photosynthetic photon flux density (Q_P), leaf-to-air vapour pressure difference (VPD), bulk leaf water potential (_x), and total hydraulic conductance (G_T) were examined in silver birch (Betula pendula Roth) with respect to leaf position in the crown. To reduce limitations caused by insufficient water supply or low light availability, experiments were also performed with branchlets cut from two different canopy layers. The intact upper-canopy leaves demonstrated 1.8–2.0 times higher (P<0.001) daily maxima of g_L compared with the lower-canopy leaves growing in the shadow of upper branches. In the morning, g_L in the shade foliage was primarily constrained by low light availability, in the afternoon, by limited water supply. Leaf conductance decreased when _x fell below certain values around midday, while the sun foliage experienced greater negative water potentials than the shade foliage. Midday stomatal openness was controlled by leaf water status and temperature, rather than by transpiration rate (E) via the feedforward mechanism. Mean G_T was 1.7 times higher (P<0.001) for the upper-canopy foliage compared to that of the lower canopy. At least 34–39% of the total resistance to the water flow from soil up to the shade foliage, and 54% up to the sun foliage, resided in 30-cm distal parts of the branches. Artificial reduction of hydraulic constraints raised _x and made g_L less sensitive to changes in both atmospheric and plant factors. Improved water supply increased g_L and E in the lower-canopy foliage, but not in the upper-canopy foliage. The results support the idea that leaves in the lower canopy are hydraulically more constrained than in the upper canopy.