Ecophysiological responses to light availability in three Blechnum species (Pteridophyta, Blechnaceae) of different ecological breadth

In Chilean evergreen temperate forest, fern species of the genus Blechnum occur in diverse microhabitats ranging from large gaps to heavily shaded understoreys. We hypothesised that differences in the ecological breadth of three co-occurring Blechnum species would be associated with differences in magnitude of ecophysiological responses to light availability. We quantified the field distribution of each species in relation to diffuse light availability (% canopy openness), and measured in situ variation in photosynthetic capacity (A), dark respiration (R _d) and specific leaf area (SLA) across the light gradient. The response of SLA of each species was also evaluated in a common garden in two light conditions (understorey and forest edge). The three Blechnum species differed significantly in the range of light environments occupied (breadth: B. chilense > B. hastatum > B. mochaenum). Despite significant interspecific differences in average A and R _d, the response of these traits to light availability did not differ among species. However, there was significant interspecific variation in both the mean value and the plasticity of SLA to light availability, the species with least ecological breadth (B. mochaenum) showing a flatter reaction norm (lower response) than its two congeners. This pattern was also found in the common garden experiment. The adjustment of leaf morphology (SLA) to light availability appears to be an important mechanism of acclimation in these Blechnum species. The narrow range of light environments occupied by B. mochaenum may be at least partly attributable to its inability to display phenotypic plasticity in SLA to changes in light availability.     

Acclimation of photosynthetic characteristics of the moss Pleurozium schreberi to among‐habitat and within‐canopy light gradients

Light availability varies strongly among moss habitats and within the moss canopy, and vertical variation in light within the canopy further interacts with the age gradient. The interacting controls by habitat and canopy light gradient and senescence have not been studied extensively. We measured light profiles, chlorophyll (Chl), carotenoid (Car) and nitrogen (N) concentrations, and photosynthetic electron transport capacity (J_max) along habitat and canopy light gradients in the widespread, temperate moss Pleurozium schreberi to separate sources of variation in moss chemical and physiological traits. We hypothesised that this species, like typical feather mosses with both apical and lateral growth, exhibits greater plasticity in the canopy than between habitats due to deeper within‐canopy light gradients. For the among‐habitat light gradient, Chl, Chl/N and Chl/Car ratio increased with decreasing light availability, indicating enhanced light harvesting in lower light and higher capacity for photoprotection in higher light. N and J_max were independent of habitat light availability. Within the upper canopy, until 50–60% above‐canopy light, changes in moss chemistry and photosynthetic characteristics were analogous to patterns observed for the between‐habitat light gradient. In contrast, deeper canopy layers reflected senescence of moss shoots, with pigment and nitrogen concentrations and photosynthetic capacity decreasing with light availability. Thus, variation in chemical and physiological traits within the moss canopy is a balance between acclimation and senescence. This study demonstrates extensive light‐dependent variation in moss photosynthetic traits, but also that between‐habitat and within‐canopy light gradient affects moss physiology and chemistry differently.     

Influence of energy flux and quality of light on the molecular organization of the photosynthetic apparatus in Scenedesmus

The photosynthetic apparatus of the unicellular green alga Scenedesmus obliquus adapts to different levels and qualities of light as documented by the fluence-rate curves of photosynthetic oxygen evolution. Cultures adapted to low fluence rates of white light (5W·m^-2) have more chlorophyll (Chl) per cell mass, a higher chlorophyll to carotenoid ratio and a doubling of the chlorophyll to cytochrome f ratio compared with cells adapted to high fluence rates of white light (20W·m^-2). Only small differences can be observed in the halfrise time of fluorescence induction, the electrophoretic profile of the pigment-protein complexes and the Chl a/Chl b-ratio. Scenedesmus cells adapted to blue light of high spectral purity demonstrate, in comparison with those adapted to red light, a higher chlorophyll content, a higher ratio of chlorophyll to carotenoid and a much higher ratio of chlorophyll to cytochrome f. Regarding these parameters and the fluence-rate curves of photosynthesis, the blue light causes the same effects as low levels of white light. In contrast, the action of red light resembles rather that of high levels of white light. Blue-light-adapted Scenedesmus cells have a smaller Chl a to Chl b ratio, a faster half-rise time of fluorescence induction and more chlorophyll in the light-harvesting system than red-light-adapted cells, as shown in the electrophoretic profile of the pigment-protein complexes. Based on these results we propose a model for the adaptation of the photosynthetic apparatus of Scenedesmus to different levels and qualities of light. In this model low as compared with high levels of white light result in an increase in the number of photosystems per electron-transport chain, but not in an increase in the size of these photosystems. The same result is obtained by adaptation to blue light. The lack of sufficient Chl b formation in red-light-adapted cells results in a decrease in the light harvesting chlorophyll-protein complexes and a photosynthetic response like that found in cells adapted to high light levels. The findings reported here confirm our earlier results in comparing blue-and red-light adaptation of the photosynthetic apparatus with adaptation to low and high levels of white light, respectively.Abbreviations Chl chlorophyll - CP chlorophyll-protein complex - DCMU 3-(3,4-dichlorophenyl)-1,1 dimethyl-urea - LHCP light harvesting chlorophyll-protein complex - LiDS lithium dodecyl sulfate - PAGE polyacrylamide gel electrophoresis - PS photosystem     

Acclimation to high irradiance in temperate deciduous trees in the field: changes in xanthophyll cycle pool size and in photosynthetic capacity along a canopy light gradient

To test the hypothesis that in temperate deciduous trees acclimation to potentially damaging high irradiances occurs via long-term adjustments in foliar photosynthetic capacity, and short-term changes in xanthophyll cycle pool size in response to weather fluctuations, nitrogen concentration and pigment composition were examined along a canopy light gradient in three species –Betula pendula, Populus tremula and Tilia cordata (from most shade intolerant to tolerant), and foliage photosynthetic potentials in P. tremula and T. cordata. Integrated quantum flux density (Q_i) incident on leaves was estimated with a method combining hemispherical photography and light measurements with quantum sensors made over the growing season. Long- and short-term light indices – average total seasonal daily integrated quantum flux density (T_s, mol m^–2 d^–1) and that of the 3 d preceding foliage sampling (T_3d) – were calculated for each sampled leaf. In addition to total integrated quantum flux density, the part of Q_i attributable to direct flux was also computed. Strong linear relationships between the capacity for photosynthetic electron transport per area (J^a_max), estimated from in situ measurements of effective quantum yield of photosystem II (PS II), and Q_i averaged over the season and over the preceding 3 d were found for all studied species. However, the major determinant of J^a_max, the product of electron transport capacity per leaf dry mass (J^m_max) and leaf dry mass per area (M_A), was M_A rather than J^m_max, which was relatively constant along the light gradient. There was evidence that J^a_max is more tightly related to T_s, which characterizes the light climate during foliar development, than to short-term integrated light, possibly because there is little flexibility in adjustments in M_A after the completion of foliar growth. Leaf chlorophyll concentrations and the investment of leaf nitrogen in chlorophyll (Chl/N) were negatively related to Q_i– an investment pattern which improves light harvesting in low light. Xanthophyll cycle pool size (VAZ, violaxanthin + antheraxanthin + zeaxanthin) either expressed per unit chlorophyll (VAZ/Chl) or as a fraction of total carotenoids (VAZ/Car) increased with increasing Q_i in all species. However, contrary to J^a_max, it tended to correlate more strongly with short-term than with long-term average integrated light. There were few interspecific differences in J^a_max, Chl/N, VAZ/Chl and VAZ/Car when the variability in light level incident to the leaves was accounted for, indicating that the foliage of both shade-intolerant and -tolerant temperate tree species possesses considerable phenotypic flexibility. Collectively these results support the view that rapid adjustment of the xanthophyll cycle pool size provides an important means for acclimation to light fluctuations in a time scale of days, during which the potential for photosynthetic quenching of excitation energy is not likely to change appreciably.     

A high proportion of blue light increases the photosynthesis capacity and leaf formation rate of Rosa × hybrida but does not affect time to flower opening

Alterations in light quality affect plant morphogenesis and photosynthetic responses but the effects vary significantly between species. Roses exhibit an irradiance‐dependent flowering control but knowledge on light quality responses is scarce. In this study we analyzed, the responses in morphology, photosynthesis and flowering of Rosa × hybrida to different blue (B) light proportions provided by light‐emitting diodes (LED, high B 20%) and high pressure sodium (HPS, low B 5%) lamps. There was a strong morphological and growth effect of the light sources but no significant difference in total dry matter production and flowering. HPS‐grown plants had significantly higher leaf area and plant height, yet a higher dry weight proportion was allocated to leaves than stems under LED. LED plants showed 20% higher photosynthetic capacity (A_max) and higher levels of soluble carbohydrates. The increase in A_max correlated with an increase in leaf mass per unit leaf area, higher stomata conductance and CO₂ exchange, total chlorophyll (Chl) content per area and Chl a/b ratio. LED‐grown leaves also displayed a more sun‐type leaf anatomy with more and longer palisade cells and a higher stomata frequency. Although floral initiation occurred at a higher leaf number in LED, the time to open flowers was the same under both light conditions. Thereby the study shows that a higher portion of B light is efficient in increasing photosynthesis performance per unit leaf area, enhancing growth and morphological changes in roses but does not affect the total Dry Matter (DM) production or time to open flower.     

Effect of foliar application of brassinolide on photosynthesis and chlorophyll fluorescence traits of <Emphasis Type="Italic">Leymus chinensis</Emphasis> under varying levels of shade

The present study was conducted to determine the effect of exogenous application of brassinolide (BR) on Leymus chinensis grown under shade, i.e., control (100% natural light), mild shade (70% natural light), and moderate shade (50% natural light). Shade substantially enhanced the plant growth, synthesis of photosynthetic pigments, photosynthetic efficiency, and chlorophyll (Chl) fluorescence attributes of L. chinensis as compared with control. The order of increase was mild shade > moderate shade > natural light except Chl content, where the order of increase was moderate shade > mild shade > natural light. Likewise, application of BR resulted in further exacerbation of plant height, plant fresh and dry mass, but less in case of Chl and carotenoids contents, gas-exchange characteristics, and Chl fluorescence attributes. The results conclude that shade significantly enhanced plant growth through alterations in physiological attributes of L. chinensis, while, application of BR may not further improve the plant growth under shade.     

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 characteristics of ornamental passion flowers grown under different light intensities

Responses of leaf gas exchange, fluorescence emission, chlorophyll concentration, and morpho-anatomical features to changes in photosynthetic photon flux density (PPFD) were studied in three wild ornamental species of Passiflora L. to select sun and shade species for landscaping projects. Artificial shade was obtained with different shading nylon nets, under field conditions, which allowed the reduction of 25, 50, and 75% of global radiation, along with a control treatment under full sunlight. For Passiflora morifolia the highest mean values of light-saturated net photosynthetic rate (P _Nmax) and light compensation point (LCP) were observed at 50 and 25% shade, respectively, while the highest values of dark respiration rate (R _D) and apparent quantum yield (α) were observed at 75% shade. For Passiflora suberosa litoralis the highest value of P _max was observed at full sunlight. The highest mean values for P _max, R _D, and LCP for Passiflora palmeri var. sublanceolata were obtained at 25% shade. The highest values of net photosynthetic rate (P _N) for P. morifolia, P. palmeri var. sublanceolata, and P. suberosa litoralis were 21.09, 16.15, and 12.36 μmol(CO₂) m⁻² s⁻¹, observed at 50 and 75% shade and full sunlight, respectively. The values of the minimal chlorophyll fluorescence (F₀) were significantly different in P. suberosa litoralis and P. palmeri var. sublanceolata, increasing with the increase of the irradiance. In contrast, the values of maximum photochemical efficiency of PSII (F_v/F_m) were significantly different only in P. suberosa litoralis, being higher at 75%, progressively reducing with the increase of PPFD levels. The total concentration of chlorophyll (Chl) was higher in shaded plants than in the ones cultivated in full sunlight. On the other hand, the values of Chl a/b ratio were reduced in shaded plants. A significant effect of shade levels on leaf area (LA) and specific leaf area (SLA) was found for the three species, whose highest mean values were observed at 75% shade. The thickness of foliar tissues was significantly higher for the three species at full sunlight and 25% shade. These results suggested that P. morifolia and P. palmeri var. sublanceolata appeared to be adapted to moderate shade conditions. P. suberosa litoralis presented higher plasticity to greater variation of the irradiance levels, while the photoinhibition was one of the limiting factors for this species at full sunlight.     

The effect of growth irradiance on leaf anatomy and photosynthesis in Acer species differing in light demand

Variation in light demand is a major factor in determining the growth and survival of trees in a forest. There is strong relation between the light‐demand and the effect of growth irradiance on leaf morphology and photosynthesis in three Acer species: A. rufinerve (light‐demanding), A. mono (intermediate) and A. palmatum (shade‐tolerant). The increase in mesophyll thickness and surface area of chloroplasts facing the intercellular airspaces (S_c) with growth irradiance was highest in A. rufinerve. Although the increase in light‐saturated photosynthesis (A_max) was similar among the species, the increase in water use efficiency (WUE) was much higher in A. rufinerve than that in the other species, indicating that the response to water limitation plays an important role in leaf photosynthetic acclimation to high light in A. rufinerve. The low CO₂ partial pressure at the carboxylation site (C_c) in A. rufinerve (130 µmol mol^?1) at high irradiance was caused by low stomatal and internal conductance to CO₂ diffusion, which minimized the increase in A_max in A. rufinerve despite its high Rubisco content. Under shade conditions, interspecific differences in leaf features were relatively small. Thus, difference in light demand related to leaf acclimation to high light rather than that to low light in the Acer species.     

Photosynthetic plasticity of two rain forest shrubs across natural gap transects

Summary Photosynthetic plasticity of two congeneric shrub species growing under natural field conditions was compared along transects spanning two canopy gaps in a Costa Rican rain forest. Piper arieianum is a shadetolerant species common in successional and mature forests, whereas P. sancti-felicis is a pioneer species abundant in abandoned clearings and large gaps. Twenty potted cuttings of each species were placed at regular intervals along two east-west transects crossing a small branch-fall gap and a large tree-fall gap. Along the transects, the percent of full sun photon flux density varied from less than 2% to 45%. After six months of growth under these conditions, leaves were monitored for incident photon flux density, photographic measures of light availability, photosynthetic capacity (A_max), leaf nitrogen content, leaf chlorophyll content, and specific leaf mass. Although both species demonstrated considerable plasticity in A_max across gap transects, P. sancti-felicis leaves had a superior capacity to track closely variation in light availability, particularly in the larger gap. For regressions of A_max on measures of light availability, P. sancti-felicis consistently showed a 3.5 to 5-fold higher coefficient of determination (R²) and a 3 to 4-fold higher slope than P. arieianum. In both species leaf nitrogen content per leaf area increased significantly with light availability, although P. sancti-felicis, again, showed a much stronger relationship between these variables. Across the transects, mean chlorophyll content per unit leaf area did not differ significantly between the species, whereas mean chlorophyll content per unit leaf dry mass was 3-times greater in leaves of P. sancti-felicis. Piper arieianum exhibited highly significant increases in chlorophyll a:b ratio with increased light availability, whereas P. sancti-felicis lacked significant variation in this trait across a gradient of light availability. Mean specific leaf mass did not vary significantly between species across the gap transects. The nature of the light acclimatory response differs quantitatively and qualitatively between these species. An important constraint on light acclimation of the shade-tolerant P. arieianum is its inability to increase photosynthetic nitrogen-use efficiency under conditions of high light availability. The lack of plasticity in chlorophyll a:b ratios does not restrict light acclimation of A_max in P. sancti-felicis. Leaves of P. arieianum exhibited symptoms of chronic photoinhibition in exposed microsites within the large gap. Species differences in the capacity to finely adjust A_max across a wide range of light conditions may be attributed to their maximum growth potential. Light acclimation in species with low maximum growth potential may be constrained at the cellular level by rates of protein and chlorophyll synthesis and at the whole-plant level by low maximum rates of uptake and supply of nutrients and water. For P. arieianum, restriction of photosynthetic plasticity is likely to limit competitive abilities of plants in high-light conditions of large gaps and clearings, whereas observed habitat restrictions for P. sancti-felicis do not appear to depend upon the highly-developed capacity for adjustment of A_max observed in this species.     

Vertical distribution of Hymenophyllaceae species among host tree microhabitats in a temperate rain forest in Southern Chile

Question: Are differences in microhabitat preferences of co‐occurring epiphytic Hymenophyllaceae species (filmy ferns) correlated with differences in ecophysiological responses to light availability and humidity in the host tree? Location: The Andean foothills in south‐central Chile. Methods: We evaluated the distribution pattern of nine filmy fern species in microhabitats that differ in light availability and humidity in four host tree species. A DCA was developed to assess Hymenophyllaceae species microhabitat preference in terms of canopy openness (CO) and relative humidity. We assessed whether differences in chlorophyll content, maximum photochemical efficiency (Fv/Fm), photosynthetic capacity (A_max), evapotranspiration (E) and instantaneous water use efficiency (WUE) are consistent with any pattern. Results: CO and relative humidity differed significantly with height in the host trees. While CO increased with height in a host tree, relative humidity decreased. DCA analysis showed that filmy fern species distribution within and among trees was mainly explained by the relative humidity of the microhabitat. Chlorophyll content, chlorophyll a/b ratio, A_max and E differed significantly among filmy fern species. A_max and E were correlated with axis 1 scores from the DCA analysis. Conclusions: The vertical distribution and abundance of filmy fern species in Chilean temperate rain forest seems to be closely related to the different microhabitats offered by host trees. This pattern may reflect interspecific differences in ecophysiological traits related both to light availability and humidity. Our results suggest that humidity is the main environmental factor driving functional responses and habitat preferences of these filmy fern species.     

Light energy partitioning in photosystems I and II during development of Nothofagus nitida growing under different light environments in the Chilean evergreen temperate rain forest

Nothofagus nitida (Phil.) Krasser (Nothofagaceae) regenerates under the shade. Nonetheless, older seedlings are commonly found at full sun. We tested the hypothesis that light capture and photochemical and non-photochemical energy dissipation of both photosystems PSI and PSII adjust with ontogeny and brighter environment. Light energy partitioning in both photosystems was studied in seedlings of different developmental stages (small 9.7 cm, tall 36 cm) under contrasting light environments (8–200 and 1,800–2,043 μmol photons m⁻² s⁻¹) in the Chilean evergreen temperate forest. Higher A _max, dark respiration, and light compensation and saturation points in sun seedlings of both developmental stages were accompanied by higher rates of electron transport. These seedlings also showed a high fraction of open PSII reaction centres and similar non-photochemical quenching at high-light in both photosystems, showing no effect of developmental stage in these parameters. Conversely, light capture, total thermal dissipation after photoinhibition, active down-regulation of antenna efficiency, and state transitions were higher in smaller seedlings than in taller ones. These changes maintain photostasis, preventing photodamage, while favouring a more oxidized quinone pool. There is an independent effect of seedling development and light acclimation on this transition from shade to sun during early ontogeny. This transition reflects short-term responses of the photosynthetic apparatus to light and longer term responses that depend on seedling developmental stage.     

Photosynthesis in chlorolichens: the influence of the habitat light regime

The hypothesis that CO₂ gas exchange and chlorophyll a fluorescence (ChlaF) of lichens vary according to the light regimes of their original habitat, as observed in vascular plants, was tested by analysing the photosynthetic performance of 12 populations of seven dorsoventral, foliose lichens collected from open, south-exposed rocks to densely shaded forests. Light response curves were induced at optimum thallus water content and ChlaF emission curves at the species-specific photon flux at which the quantum yield of CO₂ assimilation is the highest and is saturating the photosynthetic process. Photosynthetic pigments were quantified in crude extracts. The results confirm that the maximum rate of gross photosynthesis is correlated with the chlorophyll content of lichens, which is influenced by light as well as by nitrogen availability. Like leaves, shade tolerant lichens emit more ChlaF than sun-loving ones, whereas the photosynthetic quantum conversion is higher in the latter.     

Measurement of Differences in Red Chlorophyll Fluorescence and Photosynthetic Activity between Sun and Shade Leaves by Fluorescence Imaging

With a flash-lamp chlorophyll (Chl) fluorescence imaging system (FL-FIS) the photosynthetic activity of several thousand image points of intact shade and sun leaves of beech were screened in a non-destructive way within a few seconds. The photosynthetic activity was determined via imaging the Chl fluorescence at maximum F_p and steady state fluorescence F_s of the induction kinetics (Kautsky effect) and by a subsequent determination of the images of the fluorescence decrease ratio R_Fd and the ratio F_p/F_s. Both fluorescence ratios are linearly correlated to the photosynthetic CO₂ fixation rates. This imaging method permitted to detect the gradients in photosynthetic capacity and the patchiness of photosynthetic quantum conversion across the leaf. Sun leaves of beech showed a higher photosynthetic capacity and differential pigment ratios (Chl a/b and Chls/carotenoids) than shade leaves. Profile analysis and histogram of the Chl fluorescence yield and the Chl fluorescence ratios allow to quantify the differences in photosynthetic activity between different leaf parts and between sun and shade leaves with a high statistical significance.     

The energy distribution between the photosystems and light-induced changes in the stoichiometry of system I and II reaction centers in the chlorophyll b-containing alga Mantoniella squamata (Prasinophyceae)

The chlorophyll b-containing alga Mantoniella squamata was analyzed with respect to its capacity to balance the energy distribution from the light-harvesting antenna to photosystem I or photosystem II. It was shown, that this alga is unable to alter the absorption cross section of the two photosystems in terms of short-time regulations (state transitions). The energy absorbed by the LHC, which contains 60% of total photosynthetic pigments, is transferred to both photosystems without any preference. The stoichiometry of the two photosystems is found to be extremely unequal and variable during light adaptation. In high light, the molar ratio of P-680 per P-700 is found to be two, whereas under low light conditions this ratio accounts to nearly four. This very unbalanced stoichiometry of the reaction centers gives some new insights into the concept of the photosynthetic unit as well as in the importance of the regulation of the energy distribution. It is assumed that the high concentration of photosystem II can be understood as a mechanism to prevent the overexcitation of photosystem I. In addition, the changes im membrane protein pattern are not accompanied by variations in the ratio of appressed to nonappressed membranes as probed by ultrastructural analysis. It is suggested that the thylakoids are organized like a homogenous pigment bed. The lack of state transitions can be interpreted as a consequence of this unusual membrane morphology.Abbreviations Chl chlorophyll - CPa chlorophyll a-protein of PSII - CPl P-700 chlorophyll a-protein - CPD Chlorophyll packing density index - cyt f cytochrome f - FP free pigments - LHC light-harvesting complex - P_max light saturated photosynthetic rates per chlorophyll - n number of experiments - PQ plastoquinone - PS photosystem - PSU photosynthetic unit - Q_E non-photochemical quenching - Q_Q photochemical quenching     

Contrasting patterns of photosynthetic acclimation to the light environment are dependent on the differential expression of the responses to altered irradiance and spectral quality

Chl, chlorophyll
Chl a/b, ratio of chlorophyll a to chlorophyll b
Cyt f, cytochrome f
FR, far-red light
LFR, low irradiance, far-red enriched growth light
LHCII, light harvesting complex associated with PSII
LW, low irradiance, white growth light
MW, moderate irradiance, white growth light
PAR, photosynthetically active radiation
P_max, light and CO₂ saturated photosynthetic rate
PSI, photosystem I
PSII, photosystem II

Four plant species (Chamerion angustifolium, Digitalis purpurea, Brachypodium sylvaticum and Plantago lanceolata) which have previously been shown to demonstrate contrasting photosynthetic acclimatory responses to the light environment ( 33 , Plant, Cell and Environment 20, pp. 438–448) were analysed at a biochemical level. Plants were grown under low irradiance with a shade-type spectrum (LFR: 50μmol quanta m^–2 s^–1), moderately high white light (MW: 300μmol quanta m^–2 s^–1) and low irradiance white light (LW: 50μmol quanta m^–2 s^–1). The effects of light quality upon chlorophyll content and photosynthetic capacity were found to be species-dependent. A far-red dependent reduction in chlorophyll was found in three species, and an irradiance-dependent reduction was found in B. sylvaticum, which showed the greatest alteration in the xanthophyll cycle pool size of all species tested under these conditions. Chlorophyll a/b ratios were sensitive to both light quality and quantity in C. angustifolium and D. purpurea, being highest in MW, lowest in LFR, and intermediate in LW, whilst the other species showed no response. Ratios of photosystem II to photosystem I (PSII and PSI) demonstrated a strong irradiance-associated increase in all species except B. sylvaticum, whereas an increase in PSII/PSI in LFR compared to LW conditions was present in all species. A change in chlorophyll a/b was not always associated with a change in PSII/PSI, suggesting that the level of LHCII associated with each PSII varied in some species. Cytochrome f content showed an irradiance-dependent effect only, indicating a relationship with the capacity of electron transport. It is concluded that differing strategies of acclimation to the light environment demonstrated by these species results from differing strengths of expression of a series of independently regulated changes in the levels of photosynthetic components.     

Photosynthetic performance of <Emphasis Type="Italic">Lycoris radiata</Emphasis> var. <Emphasis Type="Italic">radiata</Emphasis> to shade treatments

The effects of shade on the growth, leaf photosynthetic characteristics, and chlorophyll (Chl) fluorescence parameters of Lycoris radiata var. radiata were determined under differing irradiances (15, 65, and 100% of full irradiance) within pots. The HI plants exhibited a typical decline in net photosynthetic rate (P _N) during midday, which was not observed in MI- and LI plants. This indicated a possible photoinhibition in HI plants as the ratio of variable to maximum fluorescence (F_v/F_m) value was higher and the minimal fluorescence (F₀) was lower in the, and LI plants. Diurnal patterns of stomatal conductance (g _s) and transpiration rate (E) were remarkably similar to those of P _N at each shade treatments, and the intercellular CO₂ concentration (C _i) had the opposite change trend. Under both shading conditions, the light saturation point, light compensation point and photon-saturated photosynthetic rate (P _max) became lower than those under full sunlight, and it was the opposite for the apparent quantum yield (AQY). The higher the level of shade, the lower the integrated daytime carbon gain, stomatal and epidermis cell densities, specific leaf mass (SLM), bulb mass ratio (BMR), leaf thickness, and Chl a/b ratio. In contrast, contents of Chls per dry mass (DM), leaf area ratio (LAR), leaf mass ratio (LMR), leaf length, leaf area and total leaf area per plant increased under the same shade levels to promote photon absorption and to compensate for the lower radiant energy. Therefore, when the integrated daytime carbon gain, leaf area and total leaf area per plant, which are the main factors determining the productivity of L. radiata var. radiata plant, were taken into account together, this species may be cultivated at about 60∼70% of ambient irradiance to promote its growth.     

Adaptation of epiphytic bryophytes in the understorey attributing to the correlations and trade-offs between functional traits

This study explores adaptive strategies of epiphytic bryophytes in the understorey by investigating the photosynthetic characteristics, pigment concentrations and nutrient stoichiometry, as well as other functional traits of three trunk-dwelling bryophytes in a subtropical montane cloud forest in SW China. The results showed that their light-saturated net photosynthetic rate (A_nmax?L), light saturation point (I_sat), light compensation point (I_c) and dark respiration rate (R_d) were ca 0.55, 106.72, 4.17 and 0.25?μmol?m^?2?s^?1, respectively. Furthermore, the samples demonstrated photosynthetic down-regulation under high irradiance. These photosynthetic characteristics can be explained by higher total chlorophyll concentrations, specific leaf area, chlorophyll per unit leaf N (Chl/N), lower ratio of chlorophyll a to chlorophyll b (Chl a/b) and photosynthetic nitrogen-use efficiency. We suggest that the bryophytes adapted to the shaded understorey microhabitats through a series of correlations and trade-offs between functional traits.     

Photosynthetic characteristics of dipterocarp seedlings in three tropical rain forest light environments: a basis for niche partitioning?

In three tropical rain forest light environments in Sabah, Malaysia, we compared photosynthesis in seedlings of ten climax tree species with putatively differing shade tolerances. The objectives of the study were (a) to characterise the range of photosynthetic responses in ten species of the Dipterocarpaceae and (b) to elucidate those photosynthetic characteristics that might provide a basis for niche partitioning. Seedlings were acclimated (c. 7 months) in three light environments; understorey, partial shade and a gap (140 m²). The light environments represented a gradation in median diurnal (0630–1830 hours) photon flux density (PFD) ranging from understorey (4.7 μmol m⁻² s⁻¹), through partial shade (21.2 μmol m⁻² s⁻¹) to gap (113.7 μmol m⁻² s⁻¹). Integrated diurnal PFD were in the sequence gap > partial shade > understorey (15.2, 4.7, 1.3 mol m⁻² day⁻¹, respectively). In gap-acclimated plants, species differed in the photosynthetic light-response variables apparent quantum yield, dark respiration rate, light compensation point, net saturated leaf assimilation rate (A _sat), and in stomatal conductance (g _s sat) when assimilation rate (A) was saturated. A light-demanding pioneer species (Macaranga hypoleuca) and a shade-demanding understorey species (Begonia sp.) had, respectively, higher and lower A _sat and g _s sat than the dipterocarp species. In high-light conditions A _sat and g _s sat were strongly positively correlated in dipterocarp species. Differing photosynthetic characteristics of gap-acclimated plants suggest that, in these dipterocarp species, different rates of carbon fixation may be an important factor contributing towards niche partitioning. Mean integrated diurnal A (A _diurnal) in the gap, partial shade and understory were, respectively, 122.9, 52.7, 20.5 mmol m⁻² day⁻¹. Differences occurred in A _diurnal of dipterocarp species between light environments. When Macaranga was included, differences in A _diurnal were evident in the gap and partial shade, and in both cases were attributed to the pioneer. For the variable A _diurnal, there was of a shift in the rank position of Macaranga among light environments, but a shift did not occur among the dipterocarp species. Results from this study are consistent with the idea that rates of carbon fixation per unit leaf area may contribute towards niche differentiation between the climax and single pioneer species, but not within the group of climax species. Other physiological and/or carbon allocation factors may be involved in any niche partitioning; dipterocarp species often have inherently different growth rates and susceptibility to herbivory. As an alternative to niche partitioning, dipterocarp species may co-exist in natural light environments as a result of habitat disequilibrium or purely stochastic processes. Received: 2 April 1997 / Accepted: 13 July 1997