Contribution of intercellular reflectance to photosynthesis in shade leaves

The potential contribution of intercellular light reflectance to photosynthesis was investigated by infiltrating shade leaves with mineral oil. Infiltration of leaves of Hydrophyllum canadense and Asarum canadense with mineral oil decreased adaxial leaf reflectance but increased transmittance. As a result of the large increase in transmittance, infiltration caused a decrease in absorptance of 25% and 30% at 550 and 750 nm, respectively. Thus, intercellular reflectance increased absorptance in these species by this amount. In a comparison of sun and shade leaves of Acer saccharum and Parthenocissus quinquefolia, oil infiltration decreased absorptance more in shade than in sun leaves. This difference suggests that the higher proportion of spongy mesophyll in shade leaves may increase internal light scattering and thus absorptance. The importance of the spongy mesophyll in increasing internal reflectance was also evident in comparisons of the optics of Populus leaves and in the fluorescence yield of oil-infiltrated leaves of several sun and shade species. Oil infiltration decreased the quantum yield of fluorescence (F_o) by 39–52% for shade leaves but only 21–25% for sun leaves. We conclude that the greater proportion of spongy parenchyma in shade leaves increased intercellular light scattering and thus absorptance. Direct measurements with fibre-optic light probes of the distribution of light inside leaves of Hydrophyllum canadense confirmed that oil infiltration decreased the amount of back-scattered light and that most of the light scattering for this species occurred from the middle of the palisade layer to the middle of the spongy mesophyll. We were not, however, able to assess the potential contribution of reflectance from the internal abaxial epidermis to total internal light scattering in these experiments. Using a mathematical model to compare the response of net photosynthesis (O₂, flux) to incident irradiance for control leaves of H. canadense and theoretical leaves with no intercellular reflectance, we calculated that intercellular reflectance caused a 1.97-fold increase in photosynthesis at 20 μmol m^?2s^?1 (incident photon flux density). This enhancement of absorption and photosynthesis by inter-cellular reflectance, without additional production and maintenance of photosynthetic pigments, may maintain shade leaves above the photosynthetic light compensation point between sunflecks and maintain the light induction state during protracted periods of low diffuse light.     

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.     

Distribution of UV-shielding of the epidermis of sun and shade leaves of the beech (Fagus sylvatica L.) as monitored by multi-colour fluorescence imaging

Plants can protect against damaging ultraviolet (UV) radiation by accumulating UV-absorbing substances in the epidermis of the leaves. Sun and shade leaves of a free standing beech tree (Fagus sylvatica L.) were studied for the differences in UV-shielding of the epidermis by means of multi-colour fluorescence images taken with UV and blue excitation. The distribution of the fluorescence intensity was detected over intact leaves in the emission maxima in the blue at 440 nm (F440), in the green at 520 nm (F520), in the red at 690 nm (F690) and in the far red at 740 nm (F740). Images of the logarithmic ratio between F690 excited in the blue and the UV (log (^BF690/^UVF690)) were calculated representing the relative absorption of UV in the epidermis and thus the degree of UV-shielding. It was found that UV-shielding is stronger for sun leaves than for shade leaves and better for the upper (adaxial) leaf side than for the lower (abaxial) leaf side of both leaf types. Within one leaf the highest value for the ratio log (^BF690/^UVF690) and thus the highest UV-shielding was found at the leaf rim which in broad leaves contains young tissue.     

Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light

Diurnal changes in photosynthetic gas exchange and chlorophyll fluorescence were measured under full sunlight to reveal diffusional and non‐diffusional limitations to diurnal assimilation in leaves of Arisaema heterophyllum Blume plants grown either in a riparian forest understorey (shade leaves) or in an adjacent deforested open site (sun leaves). Midday depressions of assimilation rate (A) and leaf conductance of water vapour were remarkably deeper in shade leaves than in sun leaves. To evaluate the diffusional (i.e. stomatal and leaf internal) limitation to assimilation, we used an index [1–A/A₃₅₀], in which A₃₅₀ is A at a chloroplast CO₂ concentration of 350 μ mol mol ^{? 1}. A₃₅₀ was estimated from the electron transport rate (J_T), determined fluorometrically, and the specificity factor of Rubisco (S), determined by gas exchange techniques. In sun leaves under saturating light, the index obtained after the ‘peak’ of diurnal assimilation was 70% greater than that obtained before the ‘peak’, but in shade leaves, it was only 20% greater. The photochemical efficiency of photosystem II ( Δ F/F_m ′ ) and thus J_T was considerably lower in shade leaves than in sun leaves, especially after the ‘peak’. In shade leaves but not in sun leaves, A at a photosynthetically active photon flux density (PPFD) > 500 μ mol m ^{? 2} s ^{? 1} depended positively on J_T throughout the day. Electron flows used by the carboxylation and oxygenation (J_O) of RuBP were estimated from A and J_T. In sun leaves, the J_O/J_T ratio was significantly higher after the ‘peak’, but little difference was found in shade leaves. Photorespiratory CO₂ efflux in the absence of atmospheric CO₂ was about three times higher in sun leaves than in shade leaves. We attribute the midday depression of assimilation in sun leaves to the increased rate of photorespiration caused by stomatal closure, and that in shade leaves to severe photoinhibition. Thus, for sun leaves, increased capacities for photorespiration and non‐photochemical quenching are essential to avoid photoinhibitory damage and to tolerate high leaf temperatures and water stress under excess light. The increased Rubisco content in sun leaves, which has been recognized as raising photosynthetic assimilation capacity, also contributes to increase in the capacity for photorespiration.     

High-light effects on CO2 fixation gradients across leaves

Chlorophyll fluorescence and internal patterns of ¹⁴CO₂ fixation were measured in sun and shade leaves of spinach after treatment with various light intensities. When sun leaves were irradiated with 2000μmol m^?2 s^?1 for 2h, F_V/F_M decreased by about 15%, but ¹⁴CO₂ fixation was unaffected, whereas shade leaves exhibited a 21% decrease in F_v/F_M and a 25% decrease in ¹⁴CO₂ fixation. Irradiation of sun and shade leaves with 4000μmol m^?1 for 4 h decreased F_V/F_M by 30% in sun leaves and 40% in shade leaves, while total ¹⁴CO₂ fixation decreased by 41% in sun leaves and 55% in shade leaves. After light treatment, gradients of CO₂ fixation across leaves were determined by measuring ¹⁴CO₂ fixed in paradermal leaf sections after a 10s pulse of ¹⁴CO₂. Gradients of ¹⁴CO₂ fixation in control sun and shade leaves were identified when expressed on a relative basis and normalized for leaf depth. Treatment of leaves with 2000 μmol PAR m^?2 s^?1 for 2h did not after patterns of carbon fixation across sun leaves, but slightly altered the pattern in shade leaves. In contrast, treatment of sun and shade leaves with 4000μmol m^?2 s^?1 for 4h decreased carbon fixation more in the palisade mesophyll cells than in the spongy mesophyll cells of sun and shade leaves, and fixation in medial tissue of shade leaves was dramatically decreased compared to the adaxial and abaxial tissue. The interaction between leaf anatomy and biochemical parameters involved in tolerance to photoinhibition in spinach is discussed.     

Photosynthetic costs and benefits of abaxial versus adaxial anthocyanins in Colocasia esculenta ‘Mojito’

Main conclusion

Anthocyanins in upper (adaxial) leaf tissues provide greater photoprotection than in lower (abaxial) tissues, but also predispose tissues to increased shade acclimation and, consequently, reduced photosynthetic capacity. Abaxial anthocyanins may be a compromise between these costs/benefits. Plants adapted to shaded understory environments often exhibit red/purple anthocyanin pigmentation in lower (abaxial) leaf surfaces, but rarely in upper (adaxial) surfaces. The functional significance of this color pattern in leaves is poorly understood. Here, we test the hypothesis that abaxial anthocyanins protect leaves of understory plants from photo-oxidative stress via light attenuation during periodic exposure to high incident sunlight in the forest understory, without interfering with sunlight capture and photosynthesis during shade conditions. We utilize a cultivar of Colocasia esculenta exhibiting adaxial and abaxial anthocyanin variegation within individual leaves to compare tissues with the following color patterns: green adaxial, green abaxial (GG), green adaxial, red abaxial (GR), red adaxial, green abaxial (RG), and red adaxial, red abaxial (RR). Consistent with a photoprotective function of anthocyanins, tissues exhibited symptoms of increasing photoinhibition in the order (from least to greatest): RR, RG, GR, GG. Anthocyanic tissues also showed symptoms of shade acclimation (higher total chl, lower chl a/b) in the same relative order. Inconsistent with our hypothesis, we did not observe any differences in photosynthetic CO₂ uptake under shade conditions between the tissue types. However, GG and GR had significantly (39 %) higher photosynthesis at saturating irradiance (A _sat) than RG and RR. Because tissue types did not differ in nitrogen content, these patterns likely reflect differences in resource allocation at the tissue level, with greater nitrogen allocated toward energy processing in GG and GR, and energy capture in RG and RR (consistent with relative sun/shade acclimation). We conclude that abaxial anthocyanins are likely advantageous in understory environments because they provide some photoprotection during high-light exposure, but without the cost of decreased A _sat associated with adaxial anthocyanin-induced shade syndrome.     

Inclination of sun and shade leaves influences chloroplast light harvesting and utilization

Light harvesting and utilization by chloroplasts located near the adaxial vs the abaxial surface of sun and shade leaves were examined by fluorometry in two herbaceous perennials that differed in their anatomy and leaf inclination. Leaves of Thermopsis montana had well-developed palisade and spongy mesophyll whereas the photosynthetic tissue of Smilacina stellata consisted of spongy mesophyll only. Leaf orientation depended upon the irradiance during leaf development. When grown under low-light levels, leaves of S. stellata and T. montana were nearly horizontal, whereas under high-light levels, S. stellata leaves and T. montana leaves were inclined 60⁰ and 30⁰, respectively. Leaf inclination increased the amount of light that was intercepted by the lower leaf surfaces and affected the photosynthetic properties of the chloroplasts located near the abaxial leaf surface. The slowest rates of quinone pool reduction and reoxidation were found in chloroplasts located near the adaxial leaf surface of T. montana plants grown under high light, indicating large quinone pools in these chloroplasts. Chloroplasts near the abaxial surface of low-light leaves had lower light utilization capacities as shown by photochemical quenching measurements. The amount of photosystem II (PSII) down regulation, measured from each leaf surface, was also found to be influenced by irradiance and leaf inclination. The greatest difference between down regulation monitored from the adaxial vs abaxial surfaces was found in plants with horizontal leaves. Different energy dissipation mechanisms may be employed by the two species. Values for down regulation in S. stellata were 2–3 times higher than those in T. montana, while the portion of the PSII population which was found to be Q_B nonreducing was 4–6 times lower in high light S. stellata leaves than in T. montana. All values of Stern-Volmer type nonphotochemical quenching (NPQ) from S. stellata leaves were similar when quenching analysis was performed at actinic irradiances that were higher than the irradiance to which the leaf surface was exposed during growth. In contrast, with T. montana, NPQ values from the abaxial leaf surface were up to 45% higher than those from the adaxial leaf surface regardless of growth conditions. The observed differences in chloroplast properties between species and between the adaxial and abaxial leaf surfaces may depend upon a complex interaction among light, leaf anatomy and leaf inclination.     

Origins of non-linear and dissimilar relationships between epidermal UV absorbance and UV absorbance of extracted phenolics in leaves of grapevine and barley

A recent review of climate patterns in Southern Germany has suggested significant increases in ultraviolet (UV) radiation due to decreases in cloud coverage and in cloud frequency which compound the effects of stratospheric ozone depletion. Whether such UV radiation increases result in UV damage of higher plant leaves depends partly on the capacity of UV-absorbing hydroxycinnamic acids and flavonoids located in the plant epidermis to screen out UV radiation. Epidermal UV screening is most often assessed from UV absorbance of whole-leaf extracts but in the present work, this method is critically examined. In grapevine (Vitis vinifera L.), hydroxycinnamic acid as well as mono-hydroxylated and ortho-dihydroxylated flavonoid concentrations increased in parallel with fluorometrically detected adaxial epidermal UV absorbance but only the latter class of flavonoids was associated with epidermal UV absorbance in barley (Hordeum vulgare L). For both species, curvilinear relationships between epidermal and total phenolic UV absorbance were established: initial slopes of the curves differed markedly between species. Modelling suggested that curvilinearity arises from UV-transparent epidermal areas located between vacuoles which are particularly UV-absorbing due to high levels of phenolics. The species-dependent differences were related to allocation of high amounts of phenolics in the mesophyll and abaxial epidermis in barley but not in grapevine. Both factors, optical heterogeneity and variable distribution of phenolics, severely restrict the use of phenolic absorbance to estimate true epidermal screening.     

Contribution of hydroxycinnamates and flavonoids to epidermal shielding of UV‐A and UV‐B radiation in developing rye primary leaves as assessed by ultraviolet‐induced chlorophyll fluorescence measurements

Epidermally located ultraviolet (UV)‐absorbing phenolic compounds, flavonoids and hydroxycinnamic acid esters (HCAs), can shield the underlying tissues in plants against harmful UV‐radiation. The relative importance of the two different classes of phenolic compounds for UV‐screening was a matter of recent debate. Using a non‐invasive method based on chlorophyll fluorescence measurements to estimate epidermal UV transmittance, the relationship between epidermal UV shielding and the content of the two different groups of secondary phenolic compounds in the epidermal layers and the underlying photosynthetic mesophyll of developing rye primary leaves grown under supplementary UV‐B radiation was investigated. From the fourth to the tenth day after sowing, epidermally located flavonoids increased in an age‐ and irradiation‐dependent manner, whereas mesophyll flavonoids and epidermal HCAs, mainly ferulic acid and p‐coumaric acid esters, were constitutively present and did not vary in their contents over the observed time period. There was an excellent correlation between epidermal UV‐A and UV‐B absorbances as assessed by chlorophyll fluorescence measurements and contents of epidermal flavonoids. However, HCAs showed an additional contribution to UV‐B shielding. In contrast, mesophyll flavonoids did not seem to play a respective role. When absorbances of the abaxial and adaxial epidermal layers were compared, it became apparent that in fully expanded primary leaves epidermal tissues from both sides were equally effective in absorption of UV‐radiation. However, the earlier and more UV‐exposed abaxial epidermis of young unrolling leaves showed a significantly higher absorption. It is shown that in early stages of development the epidermal HCAs are the dominant UV‐B protective compounds of the primary leaf. This function is increasingly replaced by the epidermal flavonoids during leaf development and acclimation. The application of chlorophyll fluorescence measurements has been proven to be a useful tool for estimating relative contents of these compounds in epidermal tissue.     

A molecular mechanism that confines the activity pattern of miR165 in Arabidopsis leaf primordia

In Arabidopsis leaf primordia, the expression of HD‐Zip III, which promotes tissue differentiation on the adaxial side of the leaf primordia, is repressed by miRNA165/166 (miR165/166). Small RNAs, including miRNAs, can move from cell to cell. In this study, HD‐Zip III expression was strikingly repressed by miR165/166 in the epidermis and parenchyma cells on the abaxial side of the leaf primordia compared with those on the adaxial side. We also found that the MIR165A locus, which was expressed in the abaxial epidermis, was sufficient to establish the rigid repression pattern of HD‐Zip III expression in the leaf primordia. Ectopic expression analyses of MIR165A showed that the abaxial‐biased miR165 activity in the leaf primordia was formed neither by a polarized distribution of factors affecting miR165 activity nor by a physical boundary inhibiting the cell‐to‐cell movement of miRNA between the adaxial and abaxial sides. We revealed that cis‐acting factors, including the promoter, backbone, and mature miRNA sequence of MIR165A, are necessary for the abaxial‐biased activity of miR165 in the leaf primordia. We also found that the abaxial‐determining genes YABBYs are trans‐acting factors that are necessary for the miR165 activity pattern, resulting in the rigid determination of the adaxial–abaxial boundary in leaf primordia. Thus, we proposed a molecular mechanism in which the abaxial‐biased patterning of miR165 activity is confined.     

Leaf epidermal micromorphology of Cercis (Fabaceae: Caesalpinioideae)

The micromorphology of leaf epidermal cells and stomata of all eight species and one form (11 samples) of Cercis were observed by scanning electron microscopy and light microscopy. Both the adaxial and abaxial epidermal cells are polygonal or irregular in shape; the anticlinal walls are straight and arched or undulate. Two types of stomata, which occur only on the abaxial surface of the leaves, are found in the genus. The atypical paracytic type is present in only one species, Cercis chingii, and the anomocytic type is present in all other species. Interspecific differences are minor in the genus with regard to leaf epidermal characters, except for C. chingii, which is characterized by atypical paracytic‐type stomata, a two‐lipped outer stomatal rim, the highest stomatal density and undulate and densely pitted anticlinal walls in the adaxial epidermis. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 539–547.     

Response of sun- and shade-adapted plants of Haberlea rhodopensis to desiccation

The differences in some morphological and physiological characteristics of sun- and shade-adapted Haberlea rhodopensis plants were compared. Changes in the photosynthetic activity, electrolyte leakage from leaf tissues, malondialdehyde content (MDA) and leaf anatomy were studied at different degrees of desiccation as well as after rehydration of plants. The MDA content in well-watered sun Haberlea plants was higher compared to shade plants suggesting higher lipid peroxidation, which is commonly regarded as an indicator of oxidative stress, but desiccation of plants at high light did not cause additional oxidative damage as judged by the unaffected MDA content. The electrolyte leakage from dried leaves (8% RWC) from both shade and sun plants increased fourfold indicating similar membrane damage. However, the recovery after rehydration showed that this damage was reversible. Well-watered sun plants had higher photosynthetic activity probably due to the larger thickness of the mesophyll layer in such plants. On the other hand, desiccation at high light reduced CO₂ assimilation which was in accordance with the stronger reduction of stomatal conductance. Stomata were visible only on the abaxial side of sun leaves having also higher abundance of non-glandular trichomes. Increased trichomes density and epicuticular waxes and filaments upon desiccation could help plants to increase reflection, reduce net radiation income, slow down the rate of water loss and survive adverse conditions.     

Changes of epicuticular wax induced by enhanced UV-B radiation impact on gas exchange in Brassica napus

The epicuticular wax covering on plant surface plays important roles in protecting plants against UV radiation. However, the role of epicuticular wax in affecting leaf gas exchange under enhanced ultraviolet-B (UV-B) radiation remains obscure. In the present study, different aged leaves of Brassica napus were used to analyze the responses of crystal structure and chemical constituents of epicuticular wax to UV-B radiation and the effects of such responses on gas exchange indices. Enhanced UV-B radiation significantly decreased the amount of esters in all leaves except the first leaf, amount of secondary alcohols in the second, third and fourth leaves, and amount of primary alcohols in the second and third leaves, while increased the amounts of ketones and aldehydes in the first leaf. Enhanced UV-B level had no significant effect on the amounts of alkanes and total wax in all leaves. Exposure to UV-B radiation resulted in wax fusion on adaxial leaf and stomata opening on abaxial leaf. Fusions of plates and rods on adaxial leaf surface covered most of the stomata, thereby influencing the photosynthesis in the upper mesophyll of leaves. Enhanced UV-B level significantly reduced the net photosynthesis rate (P _N) but increased the stomata conductance (g _s), concentrations of intercellular CO₂ (C _i ), and transpiration rate (E) in all leaves. Both UV-B radiation and the wax fusion induced by enhanced UV-B radiation resulted in different stomata status on abaxial and adaxial leaf surface, causing decrease of P _N, and increase of g _s, C _i and E in leaves.     

Protection by the epidermis of photosynthesis against UV-C radiation estimated by chorophyll a fluorescence

The epidermis of Argenteum mutant of Pisum sativum L. and Vicia faba L. was shown to be effective in protecting mesophyll photosynthesis from UV-C irradiation (peak 254 nm; 1.5 W m⁻²). These plants were chosen because it is easy to peel the epidermis from both sides of the leaf in Argenteum and the abaxial side in Vicia . Chlorophyll a fluorescence induction was decreased to the same extent by UV-C radiation in both leaf sides when the epidermis was removed. The fluorescence from the leaf with the epidermis inact was not affected. With irradiation of leaves with higher intensity (3.4 W m⁻²) of UV-C, the variable fluorescence was decreased by UV-C impinging on the abaxial side, but not the adaxial side in either of the plant species. A methanol extract from the mutant Pisum epidermis had a high absorbance in the UV region, and the absorbance was more than two-fold larger in adaxial epidermis than in abaxial epidermis. These results indicate that the epidermal layer, which contains substances that have high UV absorbance, protects mesophyll photosynthesis against UV radiation.     

Comparison of leaf water use efficiency of oak and sycamore in the canopy over two growing seasons

The seasonal trends in water use efficiency of sun and shade leaves of mature oak (Quercus robur) and sycamore (Acer pseudoplatanus) trees were assessed in the upper canopy of an English woodland. Intrinsic water use efficiency (net CO₂ assimilation rate/leaf conductance, A/g) was measured by gas exchange and inferred from C isotope discrimination (δ¹³C) methods. Shade leaves had consistently lower δ¹³C than sun leaves (by 1–2‰), the difference being larger in sycamore. Buds had distinct sun and shade isotopic signatures before bud break and received an influx of ¹³C-rich C before becoming net autotrophs. After leaf full expansion, δ¹³C declined by 1–2‰ gradually through the season, emphasising the importance of imported carbon in the interpretation of leaf δ¹³C values in perennial species. There was no significant difference between the two species in the value of intrinsic water use efficiency for either sun or shade leaves. For sun leaves, season-long A/g calculated from δ¹³C (72–78 μmol CO₂ [mol H₂O]⁻¹) was 10–16% higher than that obtained from gas exchange and in situ estimates of leaf boundary layer conductance. For shade leaves, the gas exchange–derived values were low, only 10–18% of the δ¹³C-derived values. This is ascribed to difficulties in obtaining a comprehensive sample of gas exchange measurements in the rapidly changing light environment.     

Size-dependent variability of leaf and shoot hydraulic conductance in silver birch

Variation in leaf and shoot hydraulic conductance was examined on detached shoots of silver birch (Betula pendula Roth), cut from the lower third (shade leaves) and upper third of the crown (sun leaves) of large trees growing in a natural temperate forest stand. Hydraulic conductances of whole shoots (K _S), leaf blades (K _lb), petioles (K _P) and branches (i.e. leafless stem; K _B) were determined by water perfusion using a high-pressure flow meter in quasi-steady state mode. The shoots were exposed to irradiance of photosynthetic photon flux density of 200–250 μmol m⁻² s⁻¹, using different light sources. K _lb depended significantly (P < 0.001) on light quality, canopy position and leaf blade area (A _L). K _lb increased from crown base to tree top, in parallel with vertical patterns of A _L. However, the analysis of data on shade and sun leaves separately revealed an opposite trend: the bigger the A _L the higher K _lb. Leaf anatomical study of birch saplings revealed that this trend is attributable to enhanced vascular development with increasing leaf area. Hydraulic traits (K _S, K _B, K _lb) of sun shoots were well co-ordinated and more strongly correlated with characteristics of shoot size than those of shade shoots, reflecting their greater evaporative load and need for stricter adjustment of hydraulic capacity with shoot size. K _S increased with increasing xylem cross-sectional area to leaf area ratio (Huber value; P < 0.01), suggesting a preferential investment in water-conducting tissue (sapwood) relative to transpiring tissue (leaves), and most likely contributing to the functional stability of the hydraulic system, essential for fast-growing pioneer species.     

葛(豆科)叶的解剖学研究

利用光学显微镜和扫描电镜观察了葛(Pueraria lobata)叶的解剖学特征。结果表明,葛叶片的上、下表皮都只有一层表皮细胞,上表皮比下表皮厚。上、下表皮都有腺毛和非腺毛。气孔主要分布在下表皮,下表皮的气孔密度为(261±17)mm-2,上表皮只有(6±3)mm-2。叶肉由两层栅栏组织细胞和一层海绵组织细胞构成。叶肉细胞中有丰富的叶绿体。在栅栏组织和海绵组织之间有一层平行于叶脉的薄壁细胞。叶脉中含有大量的草酸钙晶体。葛叶的这些形态特征与其喜阳、耐旱的特点相适应。     

Scaling sun and shade photosynthetic acclimation of Alocasia macrorrhiza to whole-plant performance – II. Simulation of carbon balance and growth at different photon flux densities

A whole-plant carbon balance model incorporating a light acclimation response was developed for Alocasia macrorrhiza based on empirical data and the current understanding of light acclimation in this species. The model was used to predict the relative growth rate (RGR) for plants that acclimated to photon flux density (PFD) by changing their leaf type, and for plants that produced only sun or shade leaves regardless of PFD. The predicted RGR was substantially higher for plants with shade leaves than for those with sun leaves at low PFD. However, the predicted RGR was not higher, and in fact was slightly lower, for plants with sun leaves than for those with shade leaves at high PFD. The decreased leaf area ratios (LARs) of the plants with sun leaves counteracted their higher photosynthetic capacities per unit leaf area (A_max). The model was manipulated by changing parameters to examine the sensitivity of RGR to variation in single factors. Overall, RGR was most sensitive to LAR and showed relatively little sensitivity to variation in A_max or maintenance respiration. Similarly, RGR was relatively insensitive to increases in leaf life-span beyond those observed. Respiration affected RGR only at low PFD, whereas A_max was moderately important only at high PFD.     

The functional ecology of shoot architecture in sun and shade plants of Heteromeles arbutifolia M. Roem., a Californian chaparral shrub

The functional roles of the contrasting morphologies of sun and shade shoots of the evergreen shrub Heteromeles arbutifolia were investigated in chaparral and understory habitats by applying a three-dimensional plant architecture simulation model, YPLANT. The simulations were shown to accurately predict the measured frequency distribution of photosynthetic photon flux density (PFD) on both the leaves and a horizontal surface in the open, and gave reasonably good agreement for the more complex light environment in the shade. The sun shoot architecture was orthotropic and characterized by steeply inclined (mean = 71^o) leaves in a spiral phyllotaxy with short internodes. This architecture resulted in relatively low light absorption efficiencies (E _A) for both diffuse and direct PFD, especially during the summer when solar elevation angles were high. Shade shoots were more plagiotropic with longer internodes and a pseudo-distichous phyllotaxis caused by bending of the petioles that positioned the leaves in a nearly horizontal plane (mean = 5^o). This shade-shoot architecture resulted in higher E _A values for both direct and diffuse PFD as compared to those of the sun shoots. Differences in E _A between sun and shade shoots and between summer and winter were related to differences in projection efficiencies as determined by leaf and solar angles, and by differences in self shading resulting from leaf overlap. The leaves exhibited photosynthetic acclimation to the sun and the shade, with the sun leaves having higher photosynthetic capacities per unit area, higher leaf mass per unit area and lower respiration rates per unit area than shade leaves. Despite having 7 times greater available PFD, sun shoots absorbed only 3 times more and had daily carbon gains only double of those of shade shoots. Simulations showed that sun and shade plants performed similarly in the open light environment, but that shade shoots substantially outperformed sun shoots in the shade light environment. The shoot architecture observed in sun plants appears to achieve an efficient compromise between maximizing carbon gain while minimizing the time that the leaf surfaces are exposed to PFDs in excess of those required for light saturation of photosynthesis and therefore potentially photoinhibitory. Received: 8 June 1997 / Accepted: 2 November 1997     

Stomatal and photosynthetic responses ofCichorium intybus leaves to sulfur dioxide treatment at different stages of plant development

Fifty-day-oldCichorium intybus Linn, plants were exposed to 1 ppm sulfur dioxide gas, 2 h per day for 7 consecutive days. Their leaves as well as those from the control plants were sampled at pre-flowering, flowering, and post-flowering stages to study their morphological, physiological, and biochemical responses to SO₂ stress. The number, dimensions, area, and biomass of leaves were less in the treated plants. Length and width of stomatal apertures on both epidermises were greater for leaves exposed to SO₂. The Stomata were longer on the adaxial epidermis, but shorter on the abaxial epidermis, except at the pre-flowering stage. Stomatal widths varied widely. Compared with the controls, the abaxial epidermis on treated leaves showed consistently lower stomatal densities as well as stomatal indices. This was also true for the adaxial epidermis during the post-flowering stage. The photosynthetic rate and stomatal conductance were reduced in the SO₂-exposed plants, but intercellular CO₂ concentrations increased at the pre-flowering stage and, subsequently, declined. Chlorophyll a, carotenoid, and total chlorophyll contents increased at the pre-flowering stage, and then decreased. The level of chlorophyllb was reduced throughout plant development compared with the untreated controls.