Shade-grown Plants Research Articles

Two Western Pacific Tabernaemontana Species Contrast in Biomass Allocation and Leaf Physiological Plasticity to Sun and Shade

The ability of plants to modify biomass allocation and leaf phenotypes to best utilize available resources has been heavily studied. No Tabernaemontana species have been included in this research agenda. Therefore, Tabernaemontana pandacaqui Poir. and Tabernaemontana rotensis (Kaneh.) B.C. Stone plants were subjected to 24% or 100% sunlight and various traits were determined to compare the relative level of plasticity. Midday net carbon dioxide assimilation was greater for T. rotensis in sun-grown plants, but greater for T. pandacaqui in shade-grown plants. Saturating light intensity and midday Fv/Fm were greater for T. rotensis in sun-grown plants, but did not differ between the species for shade-grown plants. Light compensation intensity was greater for T. rotensis than T. pandacaqui in both light treatments. Apparent quantum yield was greater for T. pandacaqui shade-grown plants but was similar for the two species in sun-grown plants. Greater relative root growth in full sun compared with shade was exhibited by T. rotensis but not by T. pandacaqui. These findings indicated that T. pandacaqui develops functional traits that improve performance in shade-grown plants, and T. rotensis develops facultative traits that enable performance in sun-grown plants. These insights into how T. pandacaqui and T. rotensis respond to sun and shade conditions add to the knowledge needed to inform the selection of niche conditions when using them in managed mixed forest plantings such as conservation and restoration sites.

Coordination of leaf photosynthetic traits and chloroplast relocation of the C4 plant Setaria palmifolia

Setaria palmifolia (Poaceae), an NADP-malic enzyme type C4 grass, is distributed from the understory to the margins of forests. To reveal how this C4 plant acclimates to habitats with contrasting light regimes, we documented the behavior of chloroplast relocation and determined patterns of chloroplast relocation and photosynthetic characteristics in response to the light of S. palmifolia grown under different light regimes. Through the observation of leaf sections and measurement of leaf transmittance (T), we confirmed that mesophyll (M) cells of S. palmifolia performed avoidance and accumulation responses, resulting in an increase and decrease in T, respectively. Measurements of gas exchange revealed that the leaf photosynthetic characteristics of S. palmifolia were modulated by growth light. Shade-grown plants significantly increased dry-weight-based chlorophyll (Chl) contents and specific leaf area (SLA), indicating increased light capture. These plants, however, displayed a reduced light-saturated photosynthetic rate (Pmax) and light saturation point (LSP) compared to unshaded plants. The patterns of chloroplast relocation were closely aligned with the photosynthetic response. Shade-grown plants displayed significantly faster kinetics, a greater extent of chloroplast relocation, and lower incident light for the induction of the avoidance response than unshaded plants. The coordination patterns of chloroplast relocation and photosynthetic response suggest that chloroplast relocation in S. palmifolia complements its leaf photosynthetic responses to light availability. The ability to adjust its photosynthetic behavior and perform chloroplast relocation may enable this C4 species to thrive in habitats with various light conditions.

Phenotypic and transcriptomic responses of the shade-grown species Panax ginseng to variable light conditions.

Elucidating how plant species respond to variable light conditions is important to understand the ecological adaptation to heterogeneous natural habitats. Plant performance and its underlying gene regulatory network have been well documented in sun-grown plants. However, the phenotypic and molecular responses of shade-grown plants under variable light conditions have remained largely unclear. We assessed the differences in phenotypic performance between Panax ginseng (shade-grown) and Arabidopsis thaliana (sun-grown) under sunlight, shade and deep-shade conditions. To further address the molecular bases underpinning the phenotypic responses, we compared time-course transcriptomic expression profiling and candidate gene structures between the two species. Our results show that, compared with arabidopsis, ginseng plants not only possess a lower degree of phenotypic plasticity among the three light conditions, but also exhibit higher photosynthetic efficiency under shade and deep-shade conditions. Further comparisons of the gene expression and structure reveal that differential transcriptional regulation together with increased copy number of photosynthesis-related genes (e.g. electron transfer and carbon fixation) may improve the photosynthetic efficiency of ginseng plants under the two shade conditions. In contrast, the inactivation of phytochrome-interacting factors (i.e. absent and no upregulation of the PIF genes) are potentially associated with the observed low degree of phenotypic plasticity of ginseng plants under variable light conditions. Our study provides new insights into how shade-grown plants respond to variable light conditions. Candidate genes related to shade adaptation in ginseng provide valuable genetic resources for future molecular breeding of high-density planting crops.

Changes in the Harpagide, Harpagoside, and Verbascoside Content of Field Grown Scrophularia lanceolata and Scrophularia marilandica in Response to Season and Shade.

Scrophularia lanceolata Pursh and Scrophularia marilandica L. are two common species within the Scrophulariaceae family that are endemic to North America. Historically, these species were used by indigenous peoples and colonialists to treat sunburn, sunstroke, frostbite, edema, as well as for blood purification, and in women’s health. Several iridoid and phenylethanoid/phenylpropanoid glycosides detected in these species, such as harpagoside and verbascoside, possess anti-inflammatory and anti-nociceptive properties. Due to the presence of anti-inflammatory metabolites and the historical uses of these species, we performed a two-year field study to determine the optimal production of these important compounds. We subjected the plants to shade treatment and analyzed differences in the metabolite composition between the two species and each of their leaves, stems, and roots at various times throughout the growing seasons. We determined that S. lanceolata plants grown in full sun produced 0.63% harpagoside per dried weight in their leaves compared to shade-grown plants (0.43%). Furthermore, S. lanceolata accumulated more harpagoside than S. marilandica (0.24%). We also found that verbascoside accumulated in the leaves of S. lanceolata and S. marilandica as the growing season progressed, while the production of this metabolite remained mostly seasonally unchanged in the roots of both species.

Comparison Between Machine Learning Models for Yield Forecast in Cocoa Crops in Santander, Colombia

The identification of influencing factors in crop yield (kg·ha-1) provides essential information for decision-making processes related to the prediction and improvement of productivity, which gives farmers the opportunity to increase their income. The current study investigates the application of multiple machine learning algorithms for cocoa yield prediction and influencing factors identification. The Support Vector Machines (SVM) and Ensemble Learning Models (Random Forests, Gradient Boosting) are compared with Least Absolute Shrinkage and Selection Operator (LASSO) regression models. The considered predictors were climate conditions, cocoa variety, fertilization level and sun exposition in an experimental crop located in Rionegro, Santander. Results showed that Gradient Boosting is the best prediction alternative with Coefficient of determination (R2) = 68%, Mean Absolute Error (MAE) = 13.32, and Root Mean Square Error (RMSE) = 20.41. The crop yield variability is explained mainly by the radiation one month before harvest, the accumulated rainfall on the harvest month, and the temperature one month before harvest. Likewise, the crop yields are evaluated based on the kind of sun exposure, and it was found that radiation one month before harvest is the most influential factor in shade-grown plants. On the other hand, rainfall and soil moisture are determining variables in sun-grown plants, which is associated with the water requirements. These results suggest a differentiated management for crops depending on the kind of sun exposure to avoid compromising productivity, since there is no significant difference in the yield of both agricultural managements.

Cyclic electron flow and light partitioning between the two photosystems in leaves of plants with different functional types

Cyclic electron flow (CEF) around photosystem I (PSI) is essential for generating additional ATP and enhancing efficient photosynthesis. Accurate estimation of CEF requires knowledge of the fractions of absorbed light by PSI (fI) and PSII (fII), which are only known for a few model species such as spinach. No measures of fI are available for C4 grasses under different irradiances. We developed a new method to estimate (1) fII in vivo by concurrently measuring linear electron flux through both photosystems left( {{text{LEF}}_{{{text{O}}_{ 2} }} } right) in leaf using membrane inlet mass spectrometry (MIMS) and total electron flux through PSII (ETR2) using chlorophyll fluorescence by a Dual-PAM at low light and (2) CEF as ETR1—{text{LEF}}_{{{text{O}}_{ 2} }}. For a C3 grass, fI was 0.5 and 0.4 under control (high light) and shade conditions, respectively. C4 species belonging to NADP-ME and NAD-ME subtypes had fI of 0.6 and PCK subtype had 0.5 under control. All shade-grown C4 species had fI of 0.6 except for NADP-ME grass which had 0.7. It was also observed that fI ranged between 0.3 and 0.5 for gymnosperm, liverwort and fern species. CEF increased with irradiance and was induced at lower irradiances in C4 grasses and fern relative to other species. CEF was greater in shade-grown plants relative to control plants except for C4 NADP-ME species. Our study reveals a range of CEF and fI values in different plant functional groups. This variation must be taken into account for improved photosynthetic calculations and modelling.

Palisade cell shape affects the light-induced chloroplast movements and leaf photosynthesis

Leaf photosynthesis is regulated by multiple factors that help the plant to adapt to fluctuating light conditions. Leaves of sun-light-grown plants are thicker and contain more columnar palisade cells than those of shade-grown plants. Light-induced chloroplast movements are also essential for efficient leaf photosynthesis and facilitate efficient light utilization in leaf cells. Previous studies have demonstrated that leaves of most of the sun-grown plants exhibited no or very weak chloroplast movements and could accomplish efficient photosynthesis under strong light. To examine the relationship between palisade cell shape, chloroplast movement and distribution, and leaf photosynthesis, we used an Arabidopsis thaliana mutant, angustifolia (an), which has thick leaves that contain columnar palisade cells similar to those in the sun-grown plants. In the highly columnar cells of an mutant leaves, chloroplast movements were restricted. Nevertheless, under white light condition (at 120 µmol m−2 s−1), the an mutant plants showed higher chlorophyll content per unit leaf area and, thus, higher light absorption by the leaves than the wild type, which resulted in enhanced photosynthesis per unit leaf area. Our findings indicate that coordinated regulation of leaf cell shape and chloroplast movement according to the light conditions is pivotal for efficient leaf photosynthesis.

Light Controls Protein Localization through Phytochrome-Mediated Alternative Promoter Selection

Alternative promoter usage is a proteome-expanding mechanism that allows multiple pre-mRNAs to be transcribed from a single gene. The impact of this mechanism on the proteome and whether it is positively exploited in normal organismal responses remain unclear. We found that the plant photoreceptor phytochrome induces genome-wide changes in alternative promoter selection in Arabidopsis thaliana. Through this mechanism, protein isoforms with different N termini are produced that display light-dependent differences in localization. For instance, shade-grown plants accumulate a cytoplasmic isoform of glycerate kinase (GLYK), an essential photorespiration enzyme that was previously thought to localize exclusively to the chloroplast. Cytoplasmic GLYK constitutes a photorespiratory bypass that alleviates fluctuating light-induced photoinhibition. Therefore, phytochrome controls alternative promoter selection to modulate protein localization in response to changing light conditions. This study suggests that alternative promoter usage represents another ubiquitous layer of gene expression regulation in eukaryotes that contributes to diversification of the proteome.

Moderate Photoinhibition of Photosystem II Protects Photosystem I from Photodamage at Chilling Stress in Tobacco Leaves.

It has been indicated that photosystem I (PSI) is susceptible to chilling-light stress in tobacco leaves, but the effect of growth light intensity on chilling-induced PSI photoinhibition in tobacco is unclear. We examined the effects of chilling temperature (4°C) associated with moderate light intensity (300 μmol photons m-2 s-1) on the activities of PSI and photosystem II (PSII) in leaves from sun- and shade-grown plants of tobacco (Nicotiana tabacum cv. k326). The sun leaves had a higher activity of alternative electron flow than the shade leaves. After 4 h chilling treatment, the sun leaves showed significantly a higher PSI photoinhibition than the shade leaves. At chilling temperature the sun leaves showed a greater electron flow from PSII to PSI, accompanying with a lower P700 oxidation ratio. When leaves were pre-treated with lincomycin, PSII activity decreased by 42% (sun leaves) and 47% (shade leaves) after 2 h exposure to the chilling-light stress, but PSI activity remained stable during the chilling-light treatment, because the electron flow from PSII to PSI was remarkably depressed. These results indicated that the stronger chilling-induced PSI photoinhibition in the sun leaves was resulted from a greater electron flow from PSII to PSI. Furthermore, moderate PSII photoinhibition depressed electron flow to PSI and then protected PSI activity against further photodamage in chilled tobacco leaves.

Sward structure and relationship between canopy height and light interception for tropical C4 grasses growing under trees

The canopy height (CH) at 95% light interception (LI) is a valuable defoliation frequency strategy used to handle variability in herbage accumulation throughout the year, mainly in C4 grasses. Such a strategy has been adopted as an open pasture management index, but defoliation frequency and intensity remain unsolved issues for shade-grown forages. A field experiment was conducted for 2 years to determine the influence of tree canopy (Eucalyptus dunnii) shading and nitrogen availability (0 and 300 kg N ha–1 year–1) on CH at 95% LI of six perennial tropical forage species. The plots were cut at 95% LI, and the height of the residual sward was kept at 50% of the corresponding CH at 95% LI. The shade level ranged from ~40% at the beginning of the experiment to ~60% at the end of summer 2013. Variations in CH at 95% LI occurred because of shading and across seasons. The range of these variations was species-dependent. Overall, species growing under trees showed higher CH, except for Paspalum notatum and Megathyrsus maximus in the first year. There was a significant increase in the length of the sheaths and leaves, as well as a decrease in tiller density and leaf : stem ratio in plants growing under trees. Nitrogen also had an impact on CH; however, its application did not compensate the shade effect on CH. Therefore, our results suggest that greater CH should be considered in case of defoliated, shade-grown plants and that such strategy might change throughout seasons.

Different roles of cyclic electron flow around photosystem I under sub-saturating and saturating light intensities in tobacco leaves.

In higher plants, the generation of proton gradient across the thylakoid membrane (ΔpH) through cyclic electron flow (CEF) has mainly two functions: (1) to generate ATP and balance the ATP/NADPH energy budget, and (2) to protect photosystems I and II against photoinhibition. The intensity of light under which plants are grown alters both CEF activity and the ATP/NADPH demand for primary metabolic processes. However, it is unclear how the role of CEF is affected by the level of irradiance that is applied during the growth and measurement periods. We studied the role of CEF at different light intensities in leaves from sun- and shade-grown plants. At 849 μmol photons m-2 s-1, both types of leaves had nearly the same degree of CEF activation. Modeling of the ATP/NADPH demand revealed that, at this light intensity, the contribution of CEF toward supplying ATP was much higher in the sun leaves. Meanwhile, the shade leaves showed higher levels of non-photochemical quenching and the P700 oxidation ratio. Therefore, at 849 μmol photons m-2 s-1, CEF mainly helped in the synthesis of ATP in the sun leaves, but functioned in photoprotection for the shade leaves. When the light intensity increased to 1976 μmol photons m-2 s-1, CEF activation was greatly enhanced in the sun leaves, but its contribution to supplying ATP changed slightly. These results indicate that the main role of CEF is altered flexibly in response to light intensity. In particular, CEF mainly contributes to balancing the ATP/NADPH energy budget under sub-saturating light intensities. When exposed to saturating light intensities, CEF mainly protects photosynthetic apparatus against photoinhibition.

Chloroplast thylakoid structure in evergreen leaves employing strong thermal energy dissipation

In nature, photosynthetic organisms cope with highly variable light environments – intensities varying over orders of magnitudes as well as rapid fluctuations over seconds-to-minutes – by alternating between (a) highly effective absorption and photochemical conversion of light levels limiting to photosynthesis and (b) powerful photoprotective thermal dissipation of potentially damaging light levels exceeding those that can be utilized in photosynthesis. Adjustments of the photosynthetic apparatus to changes in light environment involve biophysical, biochemical, and structural adjustments. We used electron micrographs to assess overall thylakoid grana structure in evergreen species that exhibit much stronger maximal levels of thermal energy dissipation than the more commonly studied annual species. Our findings indicate an association between partial or complete unstacking of thylakoid grana structure and strong reversible thermal energy dissipation that, in contrast to what has been reported for annual species with much lower maximal levels of energy dissipation, is similar to what is seen under photoinhibitory conditions. For a tropical evergreen with tall grana stacks, a loosening, or vertical unstacking, of grana was seen in sun-grown plants exhibiting pronounced pH-dependent, rapidly reversible thermal energy dissipation as well as for sudden low-to-high-light transfer of shade-grown plants that responded with photoinhibition, characterized by strong dark-sustained, pH-independent thermal energy dissipation and photosystem II (PSII) inactivation. On the other hand, full-sun exposed subalpine confers with rather short grana stacks transitioned from autumn to winter via conversion of most thylakoids from granal to stromal lamellae concomitant with photoinhibitory photosynthetic inactivation and sustained thermal energy dissipation. We propose that these two types of changes (partial or complete unstacking of grana) in thylakoid arrangement are both associated with the strong non-photochemical quenching (NPQ) of chlorophyll fluorescence (a measure of photoprotective thermal energy dissipation) unique to evergreen species rather than with PSII inactivation per se.

Species-independent down-regulation of leaf photosynthesis and respiration in response to shading: evidence from six temperate tree species.

The ability to down-regulate leaf maximum net photosynthetic capacity (Amax) and dark respiration rate (Rdark) in response to shading is thought to be an important adaptation of trees to the wide range of light environments that they are exposed to across space and time. A simple, general rule that accurately described this down-regulation would improve carbon cycle models and enhance our understanding of how forest successional diversity is maintained. In this paper, we investigated the light response of Amax and Rdark for saplings of six temperate forest tree species in New Jersey, USA, and formulated a simple model of down-regulation that could be incorporated into carbon cycle models. We found that full-sun values of Amax and Rdark differed significantly among species, but the rate of down-regulation (proportional decrease in Amax or Rdark relative to the full-sun value) in response to shade was not significantly species- or taxon-specific. Shade leaves of sun-grown plants appear to follow the same pattern of down-regulation in response to shade as leaves of shade-grown plants. Given the light level above a leaf and one species-specific number (either the full-sun Amax or full-sun Rdark), we provide a formula that can accurately predict the leaf's Amax and Rdark. We further show that most of the down regulation of per unit area Rdark and Amax is caused by reductions in leaf mass per unit area (LMA): as light decreases, leaves get thinner, while per unit mass Amax and Rdark remain approximately constant.

Photosystem II activity of typical desert plant Alhagi sparsifolia Sharp.: response to exposure to natural light after being kept in shade

Alhagi sparsifolia Sharp. is commonly considered as a type of sun plant, but shade-grown plants cannot acclimate to the strong irradiance which is normal for plant in a short time when kept in low-light conditions for a while. Alhagi sparsifolia Sharp., commonly considered as a type of sun plant, is the main vegetation found in the forelands of Taklamakan desert; this plant has an important function in wind prevention and sand fixation at the transition zone. A. sparsifolia could adapt to shaded conditions after a period of exposure to low-light conditions. However, whether or not the sun plant A. sparsifolia can adapt to natural light intensity after initial exposure to shaded conditions remains unknown. The specific light adaptation mechanism of this species is yet to be identified. In this study, the characteristics of the photosystem II (PSII) of A. sparsifolia exposed to natural light after the initial treatment under shaded lighting conditions were examined. Our results indicated that the PSII activity of A. sparsifolia under the specified condition was low; the thickness of leaves was reduced and occurred as an adaptive response to capture high amounts of light and exhibit high intensity of light-use efficiency. Shade leaves differed in terms of chlorophyll. Furthermore, the excess excitation energy has been converted to thermal dissipation energy to maintain energy balance. Shade-grown plants cannot adapt to strong irradiance, which is normal in plants for a short period when they are initially maintained under low-light conditions. Therefore, A. sparsifolia should not be considered as a typical sun plant.

Costs of defense and a test of the carbon-nutrient balance and growth-differentiation balance hypotheses for two co-occurring classes of plant defense.

One of the goals of chemical ecology is to assess costs of plant defenses. Intraspecific trade-offs between growth and defense are traditionally viewed in the context of the carbon-nutrient balance hypothesis (CNBH) and the growth-differentiation balance hypothesis (GDBH). Broadly, these hypotheses suggest that growth is limited by deficiencies in carbon or nitrogen while rates of photosynthesis remain unchanged, and the subsequent reduced growth results in the more abundant resource being invested in increased defense (mass-balance based allocation). The GDBH further predicts trade-offs in growth and defense should only be observed when resources are abundant. Most support for these hypotheses comes from work with phenolics. We examined trade-offs related to production of two classes of defenses, saponins (triterpenoids) and flavans (phenolics), in Pentaclethra macroloba (Fabaceae), an abundant tree in Costa Rican wet forests. We quantified physiological costs of plant defenses by measuring photosynthetic parameters (which are often assumed to be stable) in addition to biomass. Pentaclethra macroloba were grown in full sunlight or shade under three levels of nitrogen alone or with conspecific neighbors that could potentially alter nutrient availability via competition or facilitation. Biomass and photosynthesis were not affected by nitrogen or competition for seedlings in full sunlight, but they responded positively to nitrogen in shade-grown plants. The trade-off predicted by the GDBH between growth and metabolite production was only present between flavans and biomass in sun-grown plants (abundant resource conditions). Support was also only partial for the CNBH as flavans declined with nitrogen but saponins increased. This suggests saponin production should be considered in terms of detailed biosynthetic pathway models while phenolic production fits mass-balance based allocation models (such as the CNBH). Contrary to expectations based on the two defense hypotheses, trade-offs were found between defenses and photosynthesis, indicating that studies of plant defenses should include direct measures of physiological responses.

Phototaxis, Host Cues, and Host-Plant Finding in a Monophagous Weevil, Rhinoncomimus latipes

Rhinoncomimus latipes is a monophagous weevil used as a biological control agent for Persicaria perfoliata in the eastern United States. Density of adult R. latipes and resulting feeding damage has been shown to be higher in the sun than in the shade. This study aimed to determine whether phototaxis, sensitivity to enhanced host cues from healthier sun-grown plants, or a combination is driving this behavior by the weevil. A series of greenhouse choice tests between various combinations of plant and light conditions showed that R. latipes is positively phototactic, responsive to host cues, and preferentially attracted to sun-grown plants over shade-grown plants. From our experiments, we hypothesize two phases of dispersal and host finding in R. latipes. The initial stage is controlled primarily by phototaxis, whereas the later stage is controlled jointly by host cues and light conditions.

Differential responses of antioxidant enzymes in pioneer and late-successional tropical tree species grown under sun and shade conditions

To investigate the ability of pioneer and late-successional species to adapt to a strong light environment in a reforestation area, we examined the activities of antioxidant enzymes in relation to photosystem II, chlorophyll a fluorescence and photosynthetic pigment concentration for eight tropical tree species grown under 100% (sun) and 10% (shade) sunlight irradiation. The pioneer (early-succession) species (PS) were Cecropia pachystachya, Croton urucurana, Croton floribundus and Schinus terebinthifolius. The non-pioneer (late succession) species (LS) were Hymenaea courbaril L. var. stilbocarpa, Esenbeckia leiocarpa, Cariniana legalis and Tabebuia roseo-alba. We observed a greater decline in the ratio of variable to maximum chlorophyll a fluorescence ( F v/ F m) under full sunlight irradiation in the late-successional species than in the pioneer species. The LS species most sensitive to high irradiance were C. legalis and H. courbaril. In LS species, chlorophyll a, chlorophyll b and total chlorophyll concentrations were higher in the shade-grown plants than in plants that developed under full sunlight, but in the PS species C. floribundus and C. pachystachya, we did not observe significant changes in chlorophyll content when grown in the two contrasting environments. The carotenoids/total chlorophyll ratio increased significantly when plants developed under high-sunlight irradiation, but this response was not observed in the PS species S. terebinthifolius and C. pachystachya. The improved performance of the pioneer species in high sunlight was accompanied by an increase in superoxide dismutase (SOD, EC 1.15.1.1) activity, though no light-dependent increase in the activity of ascorbate peroxidase (APX, EC 1.11.1.11) was observed. The activity of catalase (CAT, EC 1.11.1.6) was reduced by high irradiation in both pioneer and late-successional species. Our results show that pioneer species perform better under high-sunlight irradiation than late-successional species, as indicated by increased SOD activity and a higher F v/ F m ratio. C. legalis was the LS species most susceptible to photoinhibition under full sunlight conditions. These results suggest that pioneer plants have more potential tolerance to photo-oxidative damage than late-successional species associated with the higher SOD activity found in pioneer species. Reduced photoinhibition in pioneer species probably results from their higher photosynthetic capacities, as has been observed in a previous survey carried out by our group.

Effects of coloured shade netting on the vegetative development and leaf structure of Ocimum selloi

The objective of this work was to investigate the effects of shading on the growth and development of the medicinal plant Ocimum selloi after 90 days. Seedlings were maintained under three different light treatments, namely, full sunlight vs coloured shade treatments provided by red and blue ChromatiNet® 50% coloured netting. After 90 days of cultivation, height, stem diameter, petiole length, total leaf area (TLA), specific leaf area (SLA), leaf area ratio (LAR) and leaf weight ratio (LWR) of each plant hare measured. Microscopic examination of leaf sections were conducted in order to determine the leaf blade thicknesses, together with the size and density of the stomata, the number and area of chloroplasts, and chloroplast starch grains present in the palisade parenchyma cells. Although coloured shade-grown plants were taller in comparison with those grown in full sunlight, total dry biomass together with root, stem and leaf tissue biomasses were lower. Plants exhibited phenotypic plasticity, as demonstrated by the different TLA, SLA, LAR and LWR values that were recorded under the various light treatments. Stomatal density and leaf thickness was increased in plants maintained in full sunlight owing to the expansion of the abaxial epidermis and the spongy parenchyma. Chloroplasts were more numerous and larger in plants grown under shading, whilst the accumulation of chloroplastic starch grains was greater in plants grown under red shading or in full sunlight.

Variation in photoinhibition among Sasa senanensis, Quercus mongolica, and Acer mono in the understory of a deciduous broad-leaved forest exposed to canopy gaps caused by typhoons

To elucidate mechanisms for tolerating sudden increases in light intensity following canopy gap formation, we investigated susceptibility to photoinhibition in the evergreen clonal plant bamboo, Sasa senanensis, and two deciduous broadleaf woody plants, Quercus mongolica, and Acer mono. We measured pre-dawn photochemical efficiency of photosystem II (Fv/Fm) in plants exposed to canopy gaps and in shade-grown plants through the month following gap formation. Photoinhibition (indicated by decreased Fv/Fm) was smallest in S. senanensis and largest in A. mono. S. senanensis had the highest area-based net CO2 assimilation rate (Aarea) and electron transport rate (ETR) under high light conditions. This species also had the highest leaf mass per area (LMA) and leaf nitrogen content per area (Narea). Higher values of LMA and Narea under shade conditions probably contribute to circumvent photoinhibition through maintenance of a higher ETR capacity. Q. mongolica, a gap-dependent species, had properties intermediate between S. senanensis and A. mono; it appeared less susceptible to photoinhibition than the shade-tolerant A. mono. None of the species examined had increased photosynthetic capacity 1 month after gap formation, indicating that shade-grown leaves were unable to fully acclimate to increased light.

Morphological variation of Aechmea distichantha (Bromeliaceae) in a Chaco forest: habitat and size‐related effects

Plants show different morphologies when growing in different habitats, but they also vary in their morphology with plant size. We examined differences in sun- and shade-grown plants of the bromeliad Aechmea distichantha with respect to relationships between plant size and variables related to plant architecture, biomass allocation and tank water dynamics. We selected vegetative plants from the understorey and from forest edges of a Chaco forest, encompassing the whole size range of this bromeliad. Plant biomass was positively correlated with most architectural variables and negatively correlated with most biomass allocation variables. Understorey plants were taller and had larger diameters, whereas sun plants had more leaves, larger sheath area, sheath biomass and sheath mass fraction. All tank water-related variables were positively correlated with plant biomass. Understorey plants had a greater projected leaf area, whereas sun plants had higher water content and evaporative area. Plasticity indices were higher for water-related than for allocation variables. In conclusion, there were architectural and biomass allocation differences between sun- and shade-grown plants along a size gradient, which, in turn, affected tank water-related variables.