Carotenoid composition of peridermal twigs does not fully conform to a shade acclimation hypothesis

The photosynthetic pigments of twigs in five tree and shrub species possessing chlorenchyma under a well developed, stomata-less, and highly photon absorptive periderm were analysed and compared to those of the corresponding canopy leaves. We asked whether the unavoidable shade acclimation of corticular chlorenchyma results in photosynthetic pigment complements typically found in shade leaves. As expected, chlorophyll (Chl) a/b ratios in twigs were consistently low. However, carotenoid (Car) analysis did not confirm the initial hypothesis, since twigs generally contained increased Chl-based pool sizes of the xanthophyll cycle components. The contents of photo-selective neoxanthin and lutein were high as well. Yet, -carotene content was extraordinarily low. In addition, twigs retained high pre-dawn ratios of the deepoxidized antheraxanthin and zeaxanthin, although environmental conditions were not pre-disposing for such a state. The unexpected Car composition allows the conclusion that other micro-environmental conditions within twigs (hypoxia, increased red to blue photon ratios, and extremely high CO₂ concentrations) are more important than shade in shaping the Car profiles.This revised version was published online in March 2005 with corrections to the page numbers.     

CONTRASTING PHOTOSYNTHETIC BEHAVIOR IN LEAVES AND TWIGS OF HYMENOCLEA SALSOLA,A GREEN-TWIGGED WARM DESERT SHRUB

The photosynthetic behavior of leaves and twigs was compared in Hymenoclea salsola T. and G., a subshrub of the Mohave and Sonoran deserts, in which both leaves and green twigs make substantial contributions to whole-plant carbon gain. Light saturated photosynthesis in twigs was 0.62 times that of leaves (36.9 μmol m^-2 s^-1) when plants were well watered. Similar ratios were consistently observed in contrasting the photosynthetic responses of the two organ types to light, temperature, and intercellular CO₂, regardless of whether rates were compared under saturating or highly limiting conditions of light or intercellular CO₂. These scalar differences in photosynthetic rate between leaves and green twigs under a wide range of conditions were correlated with contrasting anatomical features such as chlorenchyma volume per projected area. Under normal ambient CO₂ concentrations (350 μl 1^-1), twigs on well watered plants operated at lower intercellular CO₂ concentrations than the leaves. Possible causes of this difference are discussed with respect to performance under well-watered conditions, organ lifespans, and contrasting anatomical constraints. Twigs require larger investments than do leaves of both carbon and nitrogen per projected area of the respective organs, yet they realize lower photosynthetic rates per intercepted light. Twigs, however, fulfill additional roles besides photosynthesis such as structural support and vascular transport which does not allow them to be as anatomically specialized as leaves for photosynthesis. Twigs also have a longer expected lifespan than leaves with a larger fraction of them surviving the summer drought period. This was correlated with a greater tolerance of twig than leaf photosynthesis to low plant water potentials.     

几种树木的树皮中绿色组织的光合特性初探

测定10个树种树木上的三年生枝条树皮内绿色组织的叶绿素含量、光合速率和枝干的呼吸速率的结果表明：10个树种树皮内叶绿素含量为0.0649～0.3460mg·g^-1（FW）,叶绿素a/b为1.05～1.91,均低于叶片或针叶;树皮内绿色组织的光合速率为0.21（红松）～2.06（水曲柳）μmol（O2）·m^-2·s^-1,枝干呼吸速率1.76（红松）～3.21（落叶松）μmol（O2）·min^-1·g^-1（FW）。树皮内绿色组织的光补偿点（LCP）、光饱和点（LSP）低于叶片,而表观量子效率（AQY）高于叶片或针叶。据此认为,树皮内绿色组织适应于阴生环境,所以有较强的耐阴能力。     

Structural and functional characteristics of photosynthetic apparatus of chlorophyll-containing grape vine tissue

Photosynthesis in tissues under periderm of woody stems and shoots of perennial plants occurs in environment that is very different from the internal environment of leaf chloroplasts. These tissues are characterized by high CO₂ and low O₂ concentrations, more acidic surroundings, besides that only light which have passed through periderm reaches photosynthetic antennas. In contrast to leaves of deciduous plants chlorenchyma tissues of wintering plant organs are exposed to temperature fluctuations during all seasons, that is why the photosynthetic apparatus of woody stems has to be able to adapt to a wide range of environmental temperatures. In order to reveal unique features, which enable photosynthetic apparatus of chlorenchyma cells in woody plant organs to implement biological functions under different light and temperature conditions, we studied photosynthetic tissues of stem cortex in grapevine (Vitis vinifera L.) under normal conditions and after exposure to suboptimal temperatures and high light intensity. Comparative analysis of photosynthetic pigment composition and low-temperature chlorophyll fluorescence emission spectrum of leaves, young shoots and chlorenchyma of lignified shoots revealed relatively high level of chlorophyll b and carotenoids, and high photosystem II (PSII) to photosystem I (PSI) ratio in woody shoots. Analysis of parameters of variable chlorophyll fluorescence revealed high PSII activity in grapevine shoot cortex and demonstrated improved freeze tolerance and higher sensitivity to light of photosynthetic apparatus in grape vine in comparison to leaves. It was shown for the first time that photosynthetic apparatus in chlorenchyma cells of vine undergoes so-called “state-transition”–fast rearrangements leading to redistribution of energy between photosystems. Analysis of fatty acid (FA) compositions of lipids in examined tissues showed that the FA unsaturation index in green tissue of vine is lower than in leaves. A distinct feature of FA compositions of lipids in vine cortex was relatively high level of linoleic acid.     

Probing corticular photosynthesis through in vivo chlorophyll fluorescence measurements: evidence that high internal CO levels suppress electron flow and increase the risk of photoinhibition

Twigs of many woody plants possess chlorenchyma under a well-developed periderm which lacks stomata and impedes both gas diffusion and light penetration. The so-called corticular photosynthesis, occurring in the shade and under extremely high CO(2) concentrations, was probed in this study through in vivo chlorophyll fluorescence measurements. Field comparisons between twigs and corresponding leaves in five species indicated that both the dark- and light-adapted PSII photochemical efficiencies are considerably lower in twigs at all incident photon fluence rates, in spite of the significant attenuation of solar radiation by the periderm. Light saturation curves for linear electron transport rates (corrected according to the actual light intensities reaching twig chlorenchyma) were compatible with a shade-acclimated photosynthetic machinery, showing very low maximum electron transport rates (at approximately 5% of the corresponding leaf values) and threshold irradiances for light saturation. However, removing periderms from twig segments (i.e. relieving the twig interior form the high CO(2) partial pressures) considerably improved the light-adapted (but not the dark-adapted) PSII photochemical efficiency, allowing the assumption that the high internal CO(2) levels may interfere with the smooth functioning of photosynthesis. Indeed, laboratory experiments with twig segments equilibrated under various CO(2) levels (0.036-20%), resulted in a progressive decrease of light-adapted PSII photochemical yield, with the values obtained at 20% CO(2) being similar to those obtained with intact twigs in the field. Further experiments indicated that high CO(2) combined with high light suppressed the development of a photoprotective non-photochemical quenching through a reduction of its fast relaxing component, accompanied by a higher risk of photoinhibition. It is suggested that high internal CO(2) concentrations in twigs impede photosynthesis possibly through acidification of protoplasm and impairment of the pH-dependent high energy state quenching followed by reduction in the efficiency of heat dissipation.     

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.     

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.     

Crassulacean Acid Metabolism and Photochemical Efficiency of Photosystem II in the Adaxial and Abaxial Parts of the Succulent Leaves of Kalanchoë daigremontiana Grown at Four Photon Flux Densities

Kalanchoë daigremontiana, a species possessing crassulacean acid metabolism, was grown at four photon flux densities (1300, 400, 60, and 25 micromole photons per square meter per second). In leaves which had developed at 1300 and 400 micromole photons per square meter per second, CO₂ was mainly incorporated through the lower, shaded leaf surfaces, and the chlorenchyma adjacent to the lower surfaces showed a higher degree of nocturnal acid synthesis than the chlorenchyma adjacent to the upper surfaces. In leaves acclimated to 60 and 25 micromole photons per square meter per second, the gradient in CAM activity was reversed, i.e. more CO₂ was taken up through the upper than through the lower surfaces and nocturnal acidification was higher in the tissue next to the upper surfaces. Total net carbon gain and total nocturnal acid synthesis were highest in leaves which had developed at 400 micromole photons per square meter per second. Chlorophyll content was markedly reduced in leaves which had developed at 1300 micromole photons per square meter per second, especially in the exposed adaxial parts. There was also a sustained reduction in photosystem II photochemical efficiency as indicated by measurements of the ratio of variable over maximum chlorophyll a fluorescence. These findings suggest that, at high growth photon flux densities, the reduced activity of the exposed portions of these succulent leaves is caused by (a) the adverse effects of excess light, (b) together with a genotypic component which favors CO₂ uptake and acid synthesis in the abaxial (lower) leaf parts even when light is not or only marginally excessive. This latter component is predominant at medium photon flux densities, e.g. at 400 micromole photons per square meter per second. It becomes overridden, however, under conditions of deep shade when strongly reduced light levels in the abaxial parts of the leaf chlorenchyma severely limit photosynthesis.     

Age-dependent bark photosynthesis of aspen twigs

The photosynthetic performance of trembling aspen (Populus tremula L.) twigs and leaves was studied in relation to selected structural features of aspen bark. PFD transmittance of intact periderm was reduced by about 90% in current-year twigs through peridermal thickening. However, because of drastic changes within the bark microstructure, PFD transmittance increased in 1-year-old twig segments up to 26% of the incident PFD. On a unit surface area basis, the chlorophyll content of young twigs (425 mg Chl m^-2) almost reached that of leaves (460 mg Chl m^-2). The chlorophyll content of aspen bark chlorenchyma was clearly age-dependent, even increasing in current-year twigs with advancing internodal age. The low bark chlorophyll a/b ratios (about 2.6 compared with 3.9 in leaves) indicate that bark chloroplasts are shade-adapted. Positive net photosynthesis was not found in aspen twigs, but apparent respiration was distinctly reduced in the light due to light-driven carbon refixation (bark photosynthesis) within the chlorenchymal tissues. Under constant microclimatic conditions, dark respiration rates were strongly correlated with stem-internal CO₂ refixation. In accordance with increasing dark respiration rates, the efficiency of this carbon recycling was generally greater in the metabolically more active, younger twig segments than in older segments; carbon refixation rates reached up to 80% of dark respiration values. At least in young twigs and branches and thus in the light-exposed outer parts of tree crowns, respiratory CO₂ losses by the tree skeleton could efficiently be reduced. Refixation of carbon dioxide may be of great importance for carbon budgets in the environmentally controlled or pathogen-induced leafless states of deciduous aspen trees.     

Decline of Photosynthetic Pigments,Ribulose-1,5-Bisphosphate Carboxylase and Soluble Protein Contents,Nitrate Reductase and Photosynthetic Activities,and Changes in Thylakoid Membrane Protein Pattern in Canopy Shade Grapevine (Vitis Vinifera L. cv. Moscato Giallo) Leaves

In canopy shade leaves of grapevine (Vitis vinifera L. cv. Moscato giallo) grown in the field the contents of chlorophyll (Chl), carotenoids (Car), and soluble protein per fresh mass were lower than in sun leaves. RuBPC activity, in vivo nitrate reductase activity (indicator of nitrate utilisation), apparent electron transport rate, and photochemical fluorescence quenching were also significantly reduced in canopy shade leaves. When various photosynthetic activities were followed in isolated thylakoids, canopy shade leaves exerted a marked inhibition of whole chain and photosystem (PS) 2 activity. Smaller inhibition of PS1 activity was observed even in high-level canopy shade (HS) leaves. The artificial exogenous electron donors, DPC and NH₂OH, significantly restored the loss of PS2 activity in HS leaves. Similar results were obtained when F_v/F_m was evaluated by Chl fluorescence measurements. The marked loss of PS2 activity in canopy shade leaves was due to the loss of 47, 43, 33, 28–25, 23, 17, and 10 kDa polypeptides.     

Photosynthetic pigment contents in twigs of 24 woody species assessed by in vivo reflectance spectroscopy indicate low chlorophyll levels but high carotenoid/chlorophyll ratios

We have examined whether spectral reflectance indices used to non-destructively assess photosynthetic pigment levels and their ratios in leaves, could also be used for the same purpose in peridermal twigs. Regression lines of selected indices versus actual pigment levels, obtained from leaves and twigs of five species, suggested that semi-quantitative assessments are safe, provided that twig periderms could be easily removed. Given that, we proceeded to our next objective of screening a large number of species (24), in order to characterize their photosynthetic pigment profiles. Index comparisons between twigs and corresponding leaves indicated that twigs are characterized by lower levels of total chlorophyll and, unexpectedly, higher carotenoid/chlorophyll ratios. Moreover, the exposed and shaded sides of twigs displayed similar values for both indices in 80% of the species, suggesting that shade may not be the only factor shaping pigment levels and ratios. We discuss our results arguing that the distinct microenvironment within a twig may pose additional needs to the photosynthetic machinery, necessitating elevated carotenoid/chlorophyll ratios.     

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

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

Response of photosynthesis and chlorophyll fluorescence quenching to leaf dichotocarpism in Ligustrum vicaryi, an ornamental herb

Net photosynthetic rate of yellow upper leaves (UL) of Ligustrum vicaryi was slightly, but not significantly higher than that of green lower leaves (LL). Diurnally, maximum photochemical efficiency of photosystem 2, PS2 (F_v/F_m) of LL did not significantly decline but the UL showed fairly great daily variations. Yield of PS2 of UL showed an enantiomorphous variation to the photosynthetically active radiation and was significantly lower than in the LL. Unlike F_v/F_m, the efficiency of energy conversion in PS2 and both non-photosynthetic and photosynthetic quenching did not differ in UL and LL. Significant differences between UL and LL were found in contents of chlorophyll (Chl) a, b, and carotenoids (Car) and ratios of Chl a/b, Chl b/Chl (a+b), and Car/Chl (a+b). Leaf colour dichotocarpism in L. vicaryi was mainly caused by different photon utilization; sunflecks affected the LL.     

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     

Leaf-twig carbon isotope ratio differences in photosynthetic-twig desert shrubs

Summary Carbon isotope ratios were determined for bulk tissues of both leaves and current season twigs of 29 species of Mohave Desert shrubs. Leaf and twig tissues were found to differ in their carbon isotope ratios, only in those species which had photosynthetic twigs. These data suggest that the twigs of these species operate at lower intercellular CO₂ values than leaves, an interpretation which is consistent with available gas-exchange data. An effect of microhabitat was also evident between the mean isotope ratios of leaves from wash versus slope habitats.This research has been supported by NSF Grant BSR-8410943 and by DOE Grant DE-FGO2-86ER60399.     

Chlorophyll Fluorescence and Photosynthetic Gas Exchange Responses to Irradiance of Tree of Heaven (Ailanthus Altissima) in Contrasting Urban Environments

Sun-and shade-adapted plants of Ailanthus altissima utilized thermal-dissipative photoprotection (NPQ) across a range of photosynthetic photon flux densities (PPFD), with higher NPQ and lower maximum quantum yield of photosystem 2 photochemistry (F_v/F_m) in sun-adapted individuals, suggesting increased engagement of antennae-based quenching. Photosynthetic quantum requirements (Q_req; number of photons per CO₂) were similar in sun and shade plants, but were low and comparable to forest understory species. Diurnal measurements showed that PPFDs in both habitats were consistently above photosynthetic compensation irradiance, and frequently exceeded saturating values. In addition, sun- and shade-adapted individuals possessed stomata that tracked short-term fluctuations in PPFD. Thus A. altissima may be unique in that it couples high, shade-plant like photosynthetic efficiency with high photosynthetic capacity in high-irradiance, while stomatal attributes that optimize water use efficiency are maintained in the shade. These features may contribute to success of A. altissima in establishing in disturbance-prone urban systems, and facilitate its spread into more PPFD-limited and competitive natural ecosystems.     

Plastid terminal oxidase (PTOX) has the potential to act as a safety valve for excess excitation energy in the alpine plant species Ranunculus glacialis L.

Ranunculus glacialis leaves were tested for their plastid terminal oxidase (PTOX) content and electron flow to photorespiration and to alternative acceptors. In shade‐leaves, the PTOX and NAD(P)H dehydrogenase (NDH) content were markedly lower than in sun‐leaves. Carbon assimilation/light and C_i response curves were not different in sun‐ and shade‐leaves, but photosynthetic capacity was the highest in sun‐leaves. Based on calculation of the apparent specificity factor of ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco), the magnitude of alternative electron flow unrelated to carboxylation and oxygenation of Rubisco correlated to the PTOX content in sun‐, shade‐ and growth chamber‐leaves. Similarly, fluorescence induction kinetics indicated more complete and more rapid reoxidation of the plastoquinone (PQ) pool in sun‐ than in shade‐leaves. Blocking electron flow to assimilation, photorespiration and the Mehler reaction with appropriate inhibitors showed that sun‐leaves were able to maintain higher electron flow and PQ oxidation. The results suggest that PTOX can act as a safety valve in R. glacialis leaves under conditions where incident photon flux density (PFD) exceeds the growth PFD and under conditions where the plastoquinone pool is highly reduced. Such conditions can occur frequently in alpine climates due to rapid light and temperature changes.     

Net CO2 assimilation of taro and cocoyam as affected by shading and leaf age

Taro and cocoyam were grown outdoors in either full sun or under 40% shade. Leaves were tagged as they emerged and the effect of leaf age on net CO₂ assimilation rate (A) was determined. The effects of shading on A, transpiration (E), stomatal conductance for CO₂ (g_c) and H₂O (g_s), and water use efficiency (WUE) were also determined for leaves of a single age for each species. The effect of leaf age on A was similar for both species. Net CO₂ assimilation rates increased as leaf age increased up to 28 days with the exception of a sharp decline in A for 21 day-old leaves which corresponded to unusually low temperatures during the period of leaf expansion. A generally decreased as leaves aged beyond 28 days. Cocoyam had higher A rates than taro. Leaves of shade-grown plants had higher rates of A and E for both species at photosynthetic photon flux densities (PPFD) up to 1600 mol s^–1 m^–2. Shade-grown leaves of cocoyam had greater leaf dry weights per area (LW/A) and a trend toward higher g_c and g_s than sun-grown leaves. Shade leaves of taro had greater g_c and g₃ rates than sun-grown leaves. The data suggest that taro and cocoyam are highly adapted to moderate shade conditions.     

Effects of moderate high-temperature stress on photosynthesis in three saplings of the constructive tree species of subtropical forest

During the exposition to moderate high-temperature stress, photosynthetic rates and fluorescence of chlorophyll a were measured with a photosynthetic measurement system (Li-Cor 6400) and leaf chamber fluorometer (Li-Cor6400 LCF), respectively, in leaves of saplings, sun-adapted species (Schima superba), shade-adapted species (Cryptocarya concinna), and in mesophytic plant (Castanopsis hystrix) (42°C). The results showed that moderate high-temperature stress led to a decrease in F_v/F_>m, namely the primary photochemical quantum efficiency, indicating that moderate high-temperature stress causes a partial inhibition of PSII. It also showed that such an effect was more severe in the shade-adapted plant C. concinna than in the sun-adapted species S. superba. However, except for the sun-grown leaves of C. concinna, the moderate high-temperature stress increased the photosynthetic rate of leaves at high light intensity. Simultaneously, less photoinhibition was found to occur under high-light intensity, suggesting that the capacity of resistant-photoinhibition was stimulated by moderate high-temperature stress. The quantum yield of PSII (?_PSII) decreased in the sun-grown leaves of S. superba and C. hystrix but did not show any significant change in leaves of the shade plant C. concinna and shade-grown leaves of sun plant S. superba or the mesophytic plant C. hystrix because they already had a very low ?_PSII under this condition. Moderate high-temperature stress led to a decrease in ?_PSII/?_CO2 ratios, an estimate of the quantum requirement for CO2 assimilation, in the sun plant S. superba and the mesophytic plant C. hystrix because they were associated with the dissipation of a lower fraction of excitation energy. However, no significant changes were found in shade plant C. concinna and in shade-grown leaves of the other examined plants. The effect of moderate high-temperature (42°C) on photosynthesis depends on species and leaf type (sun and shade leaves) in the saplings of subtropical broad-leaved trees.     

On the role of orientation in reducing photoinhibitory damage in photosynthetic-twig desert shrubs

Quantum yields for O₂ evolution were measured in photosynthetic twigs and leaves of Hymenoclea salsola and Senecio douglasii, two common shrubs of the deserts of western North America. When exposed to long-term drought, quantum yields of leaves and twigs remained constant up to the point of leaf abscision in both species. As water stress developed further, quantum yields began to decline in twigs; the extent of this quantum yield reduction was dependent on incident photon flux density. The reduction in quantum yield in twigs, which have a near-vertical orientation, was greatly accelerated when twigs were reoriented to the near-horizontal inclination typical of leaves. The reductions in quantum yield were not rapidly reversible and are interpreted as indicating photoinhibitory damage. The results are discussed in terms of the role that a near-vertical orientation might serve in maintaining photosynthetic structures through a drought period.