Chlorophyll fluorescence and photosynthetic response to light in 1-year-old needles during spring and early summer in <Emphasis Type="Italic">Pinus leucodermis</Emphasis>

Ten-year-old trees from four Italian populations of Pinus leucodermis (populations A, B, C and D), which were collected from different sites at different altitudes, were grown near Florence, Italy. Needle CO₂ gas exchange and chlorophyll fluorescence response to increasing light intensities were evaluated; gas exchange and chlorophyll fluorescence variation between April and July were also monitored. Populations A, B and C showed a similar photosynthetic response to increasing photosynthetic photon flux density (PPFD) intensities, while at various light intensities population D, which originated from the highest altitude, showed the highest photosynthetic rates. In this population photosynthesis was saturated at PPFDs higher than 900 µmol m^-2s^-1 and a slow decrease of effective photosystem II quantum yield and F'_V/F'_M in response to increasing PPFDs were found. The same trees also showed a faster recovery in photosynthesis from limitations induced by winter temperatures than the other three populations. This work showed that photosynthetic response to light in population D was different from the other populations; trees from this population were probably naturally selected to prevent photoinhibition due to excess light.     

Photosynthetic sunfleck utilization potential of understory saplings growing under elevated CO<Subscript>2</Subscript> in FACE

Few studies have evaluated elevated CO₂ responses of trees in variable light despite its prevalence in forest understories and its potential importance for sapling survival. We studied two shade-tolerant species (Acer rubrum, Cornus florida) and two shade-intolerant species (Liquidambar styraciflua, Liriodendron tulipifera) growing in the understory of a Pinus taeda plantation under ambient and ambient+200 ppm CO₂ in a free air carbon enrichment (FACE) experiment. Photosynthetic and stomatal responses to artificial changes in light intensity were measured on saplings to determine rates of induction gain under saturating light and induction loss under shade. We expected that growth in elevated CO₂ would alter photosynthetic responses to variable light in these understory saplings. The results showed that elevated CO₂ caused the expected enhancement in steady-state photosynthesis in both high and low light, but did not affect overall stomatal conductance or rates of induction gain in the four species. Induction loss after relatively short shade periods (<6 min) was slower in trees grown in elevated CO₂ than in trees grown in ambient CO₂ despite similar decreases in stomatal conductance. As a result leaves grown in elevated CO₂ that maintained induction well in shade had higher carbon gain during subsequent light flecks than was expected from steady-state light response measurements. Thus, when frequent sunflecks maintain stomatal conductance and photosynthetic induction during the day, enhancements of long-term carbon gain by elevated CO₂ could be underestimated by steady-state photosynthetic measures. With respect to species differences, both a tolerant, A. rubrum, and an intolerant species, L. tulipifera, showed rapid induction gain, but A. rubrum also lost induction rapidly (c. 12 min) in shade. These results, as well as those from independent studies in the literature, show that induction dynamics are not closely related to species shade tolerance. Therefore, it cannot be concluded that shade-tolerant species necessarily induce faster in the variable light conditions common in understories. Although our study is the first to examine dynamic photosynthetic responses to variable light in contrasting species in elevated CO₂, studies on ecologically diverse species will be required to establish whether shade-tolerant and -intolerant species show different photosynthetic responses in elevated CO₂ during sunflecks. We conclude that elevated CO₂ affects dynamic gas exchange most strongly via photosynthetic enhancement during induction as well as in the steady state. Received: 1 April 1999 / Accepted: 16 August 1999     

The greater seedling high-light tolerance of <Emphasis Type="Italic">Quercus robur</Emphasis> over <Emphasis Type="Italic">Fagus sylvatica </Emphasis>is linked to a greater physiological plasticity

The responses of Quercus robur (oak) and Fagus sylvatica (beech) seedlings to four different light environments (full, 50%, 40% and 15% sunlight) and to a rapid increase in irradiance were explored during the summer, after 2 years of growth in a forest nursery at Nancy (France). Significant differences between the two species were found for most variables. Phenotypic plasticity for morphological variables (root-shoot ratio, leaf size, leaf weight ratio) was higher in beech than in oak, while the reverse was true for anatomical (stomatal density, epidermis thickness, exchange surface area of the palisade parenchyma) and physiological (maximum photosynthetic rate, stomatal conductance, Rubisco activity) variables. Predawn photochemical efficiency (F_v/F_m) was higher in oak than in beech in all light environments except in 15% sunlight. F_v/F_m was significantly lower in 100% sunlight than in the other light environments in beech but not in oak. Maximum photosynthetic rates (A_max) increased with increasing light availability in the two species but they were always higher in oak than in beech. Oak exhibited higher Rubisco activity than beech in full sunlight. The transfer of shade-adapted seedlings to the open caused a decrease of F_v/F_m, which was larger for beech than for oak. Transferred oak but not beech plants recovered gradually to the control F_v/F_m values. The decreased chlorophyll content and the increased non-photochemical quenching observed in high-light beech seedlings were not enough to avoid photoinhibition. The results suggest that a greater tolerance of strong irradiance is linked to an enhanced physiological plasticity (variables related to photosynthesis), while shade tolerance relies on an enhanced plasticity in light-harvesting variables (crown morphology and chlorophyll content).     

Modeling dynamic understory photosynthesis of contrasting species in ambient and elevated carbon dioxide

Dynamic responses of understory plants to sunflecks have been extensively studied, but how much differences in dynamic light responses affect daily photosynthesis (A_day) is still the subject of active research. Recent models of dynamic photosynthesis have provided a quantitative tool that allows the critical assessment of the importance of these sunfleck responses on A_day. Here we used a dynamic photosynthesis model to assess differences in four species that were growing in ambient and elevated CO₂. We hypothesized that Liriodendron tulipifera, a species with rapid photosynthetic induction gain and slow induction loss, would have the least limitations to sunfleck photosynthesis relative to the other three species (Acer rubrum, Cornus florida, Liquidambar styraciflua). As a consequence, L. tulipifera should have the highest A_day in an understory environment, despite being the least shade tolerant of the species tested. We further hypothesized that daily photosynthetic enhancement by elevated CO₂ would differ from enhancement levels observed during light-saturated, steady-state measurements. Both hypotheses were supported by the model results under conditions of low daily photosynthetic photon flux density (PFD; <3% of the above-canopy PFD). However, under moderate PFD (10-20% of the above-canopy PFD), differences in dynamic sunfleck responses had no direct impact on A_day for any of the species, since stomatal and photosynthetic induction limitations to sunfleck photosynthesis were small. Thus, the relative species ranking in A_day under moderate PFD closely matched their rankings in steady-state measurements of light-saturated photosynthesis. Similarly, under elevated CO₂, enhancement of modeled A_day over A_day at ambient CO₂ matched the enhancement measured under light saturation. Thus, the effects of species-specific differences in dynamic sunfleck responses, and differences in elevated CO₂ responses of daily photosynthesis, are most important in marginal light environments.     

Relationship between thylakoid electron transport and photosynthetic CO2 uptake in leaves of three maize (Zea mays L.) hybrids

The introduction of a more efficient means of measuring leaf photosynthetic rates under field conditions may help to clarify the relationship between single leaf photosynthesis and crop growth rates of commercial maize hybrids. A large body of evidence suggests that gross photosynthesis (A_G) of maize leaves can be accurately estimated from measurements of thylakoid electron transport rates (ETR) using chlorophyll fluorescence techniques. However, before this technique can be adopted, it will first be necessary to determine how the relationship between chlorophyll fluorescence and CO2 assimilation is affected by the non-steady state PPFD conditions which predominate in the field. Also, it must be determined if the relationship is stable across different maize genotypes, and across phenological stages. In the present work, the relationship between ETR and A_G was examined in leaves of three maize hybrids by making simultaneous measurements of leaf gas exchange and chlorophyll fluorescence, both under controlled environment conditions and in the field. Under steady-state conditions, a linear relationship between ETR and A_G was observed, although a slight deviation from linearity was apparent at low A_G. This deviation may arise from an error in the assumption that respiration in illuminated leaves is equivalent to respiration in darkened leaves. The relationship between chlorophyll fluorescence and photosynthetic CO2 assimilation was not stable during fluctuations in incident PPFD. Since even minor (e.g. 20%) fluctuations in incident PPFD can produce sustained ( > 20 s) departures from the mean relationship between ETR and A_G, chlorophyll fluorometry can only provide an accurate estimate of actual CO2 assimilation rates under relatively stable PPFD conditions. In the field, the mean value of ETR / A_G during the early part of the season (4.70 ± 0.07) was very similar to that observed in indoor-grown plants in the vegetative stage (4.60 ± 0.09); however, ETR / A_G increased significantly over the growing season, reaching 5.00 ± 0.09 by the late grain-filling stage. Differences in ETR / A_G among the three genotypes examined were small (less than 1% of the mean) and not statistically significant, suggesting that chlorophyll fluorometry can be used as the basis of a fair comparison of leaf photosynthetic rates among different maize cultivars.     

Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N_area) were determined for individuals, and leaf photosynthesis was measured as a function of N_area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P_mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P_mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P_mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P_mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N_area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N_area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.     

The influence of UV-B radiation on light-dependent photosynthetic performance in <Emphasis Type="Italic">Sanionia uncinata </Emphasis>(Hedw.) Loeske in Antarctica

Photosynthetic activity of the moss Sanionia uncinata (Hedw.) Loeske was investigated on Léonie Island (67°35'S, 68°20'W, Antarctic Peninsula) in response to short-term changes of UV-B radiation. The UV-environment of natural mat formations dominated by S. uncinata was altered using filter screens. Two filter experiments were conducted in the Antarctic summers 1998 and 1999. A third filter experiment was conducted during springtime ozone depletion in October 1998. Photosynthetic activity of S. uncinata was mainly determined by photosynthetically active photon flux density (PPFD). Light response of relative electron transport rate through photosystem II (rel ETR=jF/Fm'2PPFD) remained unaffected by ambient summer levels of UV-B radiation. The same was found for net photosynthesis and dark respiration. In October 1998, S. uncinata was mainly metabolically inactive due to low temperatures. No significant levels of DNA-damage measured as cyclobutane pyrimidine dimers (CPDs) were induced by ambient summer levels of UV-B. Artificially enhanced UV-B radiation supplying a Setlow-DNA-dose of 8.7 kJ m^ф day^у UV-B led to formation of 7Dž CPD (10⁶ nucleotides)^-1. It is concluded that current ambient summer levels of UV-B radiation do not affect photosynthetic activity in S. uncinata.     

Photosynthetic induction in saplings of three shade-tolerant tree species: comparing understorey and gap habitats in a French Guiana rain forest

The photosynthetic induction response under constant and fluctuating light was examined in naturally occurring saplings (about 0.5-2 m in height) of three shade-tolerant tree species, Pourouma bicolor spp digitata, Dicorynia guianensis, and Vouacapoua americana, growing in bright gaps and in the shaded understorey in a Neotropical rain forest. Light availability to saplings was estimated by hemispherical photography. Photosynthetic induction was measured in the morning on leaves that had not yet experienced direct sunlight. In Dicorynia, the maximum net photosynthesis rate (A_max) was similar between forest environments (ca 4 µmol m^-2 s^-1), whereas for the two other species, it was twice as high in gaps (ca 7.5) as in the understorey (ca 4.5). However, the time required to reach 90% of A_max did not differ among species, and was short, 7-11 min. Biochemical induction was fast in leaves of Pourouma, as about 3 min were needed to reach 75% of maximum carboxylation capacity (V_cmax); the two other species needed 4-5 min. When induction continued after reaching 75% of V_cmax, stomatal conductance increased in Pourouma only (ca 80%), causing a further increase in its net photosynthesis rate. When fully induced leaves were shaded for 20 min, loss of induction was moderate in all species. However, gap saplings of Dicorynia had a rapid induction loss (ca 80%), which was mainly due to biochemical limitation as stomatal conductance decreased only slowly. When leaves were exposed to a series of lightflecks separated by short periods of low light, photosynthetic induction increased substantially and to a similar extent in all species. Although A_max was much lower in old than in young leaves as measured in Dicorynia and Vouacapoua, variables of the dynamic response of photosynthesis to a change in light tended to be similar between young and old leaves. Old leaves, therefore, might remain important for whole-plant carbon gain, especially in understorey environments. The three shade-tolerant species show that, particularly in low light, they are capable of efficient sunfleck utilization.     

Gas exchange and chlorophyll fluorescence responses of <Emphasis Type="Italic">Pinus radiata</Emphasis> D. Don seedlings during and after several storage regimes and their effects on post-planting survival

Post-storage gas exchange parameters like CO₂ assimilation, stomatal conductance, transpiration, water use efficiency and intercellular CO₂ concentrations, together with several chlorophyll a fluorescence parameters: F_o, F_v, F_v/F_m, F_m/F_o and F_v/F_o were examined in radiata pine (Pinus radiata D. Don) seedlings that were stored for 1, 8 or 15 days at 4° or 10°C with or without soil around the roots. Results were analysed in relation to post-storage water potential and electrolyte leakage in order to forecast their vitality (root growth potential) following cold storage, and post-planting survival potential under optimal conditions. During storage at 4° and 10°C, photosynthesis was reduced, being more pronounced in bare-root seedlings than in seedlings with soil around the roots. The depletion of CO₂ assimilation seemed not to be solely a stomatal effect as effects on chloroplasts contributed to this photosynthetic inhibition. Thus, the fall in the ratios F_v/F_m, F_v/F_o and F_m/F_o indicated photochemical apparatus damage during storage. Photosynthetic rate was positively correlated with the root growth index and new root length showing that new root growth is dependent primarily on current photosynthesis. Pre-planting exposure of bare-root radiata pine seedlings to temperatures of 10°C for more than 24 h during transportation or storage is not recommended.     

Effects of elevated CO<Subscript>2</Subscript> on foliar chemistry of saplings of nine species of tropical tree

This study examined the effects of elevated CO₂ on secondary metabolites for saplings of tropical trees. In the first experiment, nine species of trees were grown in the ground in open-top chambers in central Panama at ambient and elevated CO₂ (about twice ambient). On average, leaf phenolic contents were 48% higher under elevated CO₂. Biomass accumulation was not affected by CO₂, but starch, total non-structural carbohydrates and C/N ratios all increased. In a second experiment with Ficus, an early successional species, and Virola, a late successional species, treatments were enriched for both CO₂ and nutrients. For both species, nutrient fertilization increased plant growth and decreased leaf carbohydrates, C/N ratios and phenolic contents, as predicted by the carbon/nutrient balance hypothesis. Changes in leaf C/N levels were correlated with changes in phenolic contents for Virola (r=0.95, P<0.05), but not for Ficus. Thus, elevated CO₂, particularly under conditions of low soil fertility, significantly increased phenolic content as well as the C/N ratio of leaves. The magnitude of the changes is sufficient to negatively affect herbivore growth, survival and fecundity, which should have impacts on plant/herbivore interactions.     

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.     

Differential impacts of long-term (CO<Subscript>2</Subscript>) and O<Subscript>3</Subscript> exposure on growth of northern conifer and deciduous tree species

The long-term effects of elevated CO₂ and CO₂+O₃ concentrations on the growth allocation in northern provenances of Norway spruce [Picea abies (L.) Karst.], Scots pine [Pinus sylvestris (L.)] and pubescent birch clones (Betula pubescens Ehrh.) were examined in open-top chambers after a 4-year-long experiment. The total biomass responses of the tree seedlings to increased CO₂ and CO₂+O₃ concentrations were not statistically significant and varied between the provenances and species. The seedlings of northern origin were the least sensitive in their response to treatments. The total biomass of the Norway spruce seedlings slightly decreased in response to CO₂ in three provenances. Scots pine from the local provenance had a slight biomass increase after elevated CO₂+O₃ treatment. The slower-growing birch clone seemed to benefit from elevated CO₂, whereas in the faster-growing clone, reductions in biomass accumulation were seen. The combined CO₂+O₃ treatment reduced the positive effects of elevated CO₂, especially in the slower-growing birches. Observations of significant effects were limited to a few parameters. Carbon dioxide treatment decreased needle dry weight of Norway spruce in one northern provenance. The needle and wood dry weight increased (CO₂ + O₃) in local Scots pine. Significant birch response was limited to increased fine root density (O₃ + CO₂) in the inland clone. The diverse effects of elevated CO₂ and CO₂ +O₃ on seedling growth and biomass provide evidence that exposure of northern trees to the enhanced variable CO₂ and O₃ concentrations of the future will have varied effects on the growth of these species. The direction and magnitude of those effects will differ depending on species and origins.     

Leaves of Japanese oak (Quercus mongolica var. crispula) mitigate photoinhibition by adjusting electron transport capacities and thermal energy dissipation along the intra-canopy light gradient

We investigated the morphological and physiological acclimation of leaves grown within a canopy of Japanese oak tree (Quercus mongolica var. crispula) in terms of the susceptibility to photoinhibition under various growth light conditions. The maximum rates of photosynthesis (P(max) ) and electron transport (ETR(max) ) were higher in mature leaves grown under stronger light with higher area-based leaf nitrogen (N) content closely associated with higher leaf mass per area. The net photosynthetic (P(n) ) and electron transport (ETR) rates corresponding to the daily peak photosynthetic photon flux density (PPFD(max) ) during leaf maturation were almost comparable to P(max) and ETR(max) , respectively. Conversely, P(n) and ETR at the daily average PPFD (PPFD(avg) ) were substantially low in shade-grown leaves when compared with P(max) and ETR(max) . The susceptibility to photoinhibition at PPFD(max) , i.e. at sunflecks for the shade-grown leaves, was assessed by the rate of excess energy production. Although sun leaves showed higher rates of electron transport and thermal energy dissipation than shade leaves under PPFD(max) conditions, the rate of excess energy production was almost constant across shade to sun leaves. The shade leaves of the Japanese oak grown within a crown were suggested to adjust their N investment to maintain higher photosynthetic capacities compared with those required to maximize the net carbon gain, which may facilitate the dissipation of the excessive light energy of sunflecks to circumvent photoinhibition in cooperation with thermal energy dissipation.     

Photosynthetic induction and leaf carbon gain in the tropical understorey epiphyte,Aspasia principissa

Gas exchange of the understorey epiphyte Aspasia principissa was studied in fluctuating light conditions both in the laboratory and in the field, testing the hypothesis that vascular epiphytes differ from most terrestrial understorey plants in showing a higher priority for water conservation. Consequently, a slow response of stomatal conductance to sudden increases in incident photon flux density (PFD) was expected, as was a fast loss of induction after such a light fleck. Results were only partly consistent with these expectations. Full induction of photosynthesis was indeed very slow and was not reached before, respectively, 40 and 60 min of saturating PFD in the field and the laboratory. In contrast, kinetics of induction loss were comparable with those of most terrestrial species studied to date. The overall impact of light flecks on in situ carbon gain again fulfilled expectations, being rather limited: the observed carbon gain was only approx. 66% of the potential carbon gain estimated from a square-wave response model. It is concluded that in the drought-prone epiphytic habitat of a moist lowland forest, water conservation takes priority over carbon gain, which severely limits the use of light flecks for CO(2) fixation in vascular epiphytes.     

The photobiont determines the pattern of photosynthetic activity within a single lichen thallus containing cyanobacterial and green algal sectors (photosymbiodeme)

Photosystem activity status of the green algal (Pseudocyphellaria lividofusca) and cyanobacterial (P. knightii) components of a photosymbiodeme were continuously monitored in the field over a period of 35 days. The photosymbiodeme grew on a Nothofagus menziesii tree at Lake Waikaremoana, Urewera National Park, North Island, New Zealand. Two Mini-PAM fluorometers were placed so that the chlorophyll a fluorescence, temperature and PPFD (photosynthetically active photon flux density) could be recorded every 30 min for green algal and cyanobacterial parts of the thallus. Microclimate conditions were also recorded with a datalogger. The study confirmed the already known ability of green algal lichens to reactivate from high humidity alone whilst cyanobacterial species need liquid water, here obtained from rainfall. The photosystems of P. lividofusca were activated on every day and positive ETR (relative electron transport rate) occurred on all but 3 days. Activation level depended on the overnight relative humidity. P. knightii was activated and had positive ETR on only 13 days when rainfall had occurred. Both species were mostly inactive above 12°C but differed at low temperatures. P. knightii showed no activation at very low temperatures, -2 to 0°C, since these only occurred on clear, rain-free nights. PPFD was always very low, mostly below 80 µmol m^-2 s^-1, and both species were inactive at higher PPFD. The three-dimensional structure of the thallus seemed to contribute to the hydration. The cyanobacterial sectors were more appressed to the trunk and needed substantial rainfall to rewet whereas the green algal lobes were more distant from the trunk and this probably caused more rapid desiccation as well as lower temperatures. It is suggested that the longer active periods for photosynthesis by P. lividofusca are balanced by several factors: first, depressed net photosynthesis at high thallus water contents after rainfall, a feature not shown by P. knightii; second, possible lower maximal net photosynthetic rates; and third, the possibility of greater respiratory rates when thalli have been hydrated by high relative humidity. There is little evidence for high PPFD differently affecting the photosymbiodeme components since sustained, high PPFD did not occur. It has been reported that the photosystems of cyanobacterial species from photosymbiodemes can reactivate at high relative humidity but the results obtained here suggest that it is not ecologically significant.     

Responses of leaf photosynthesis,pigments and chlorophyll fluorescence within canopy position in a boreal grass (<Emphasis Type="Italic">Phalaris arundinacea</Emphasis> L.) to elevated temperature and CO<Subscript>2</Subscript> under varying water regimes

The effects of elevated growth temperature (ambient + 3.5°C) and CO₂ (700 μmol mol⁻¹) on leaf photosynthesis, pigments and chlorophyll fluorescence of a boreal perennial grass (Phalaris arundinacea L.) under different water regimes (well watered to water shortage) were investigated. Layer-specific measurements were conducted on the top (younger leaf) and low (older leaf) canopy positions of the plants after anthesis. During the early development stages, elevated temperature enhanced the maximum rate of photosynthesis (P _max) of the top layer leaves and the aboveground biomass, which resulted in earlier senescence and lower photosynthesis and biomass at the later periods. At the stage of plant maturity, the content of chlorophyll (Chl), leaf nitrogen (N_L), and light response of effective photochemical efficiency (Φ_PSII) and electron transport rate (ETR) was significantly lower under elevated temperature than ambient temperature in leaves at both layers. CO₂ enrichment enhanced the photosynthesis but led to a decline of N_L and Chl content, as well as lower fluorescence parameters of Φ_PSII and ETR in leaves at both layers. In addition, the down-regulation by CO₂ elevation was significant at the low canopy position. Regardless of climate treatment, the water shortage had a strongly negative effect on the photosynthesis, biomass growth, and fluorescence parameters, particularly in the leaves from the low canopy position. Elevated temperature exacerbated the impact of water shortage, while CO₂ enrichment slightly alleviated the drought-induced adverse effects on P _max. We suggest that the light response of Φ_PSII and ETR, being more sensitive to leaf-age classes, reflect the photosynthetic responses to climatic treatments and drought stress better than the fluorescence parameters under dark adaptation.     

Reduction of the primary donor P700 of photosystem I during steady-state photosynthesis under low light in <Emphasis Type="Italic">Arabidopsis</Emphasis>

During steady-state photosynthesis in low-light, 830-nm absorption (A₈₃₀) by leaves was close to that in darkness in Arabidopsis, indicating that the primary donor P700 in the reaction center of photosystem I (PSI) was in reduced form. However, P700 was not fully oxidized by a saturating light pulse, suggesting the presence of a population of PSI centers with reduced P700 that remains thermodynamically stable during the application of the saturating light pulse (i.e., reduced-inactive P700). To substantiate this, the effects of methyl viologen (MV) and far-red light on P700 oxidation by the saturating light pulse were analyzed, and the cumulative effects of repetitive application of the saturating light pulse on photosynthesis were analyzed using a mutant crr2-2 with impaired PSI cyclic electron flow. We concluded that the reduced-inactive P700 in low-light as revealed by saturating light pulse indicates limitations of electron flow at the PSI acceptor side.     

Cytological investigation of male sterility in sorghum caused by a dominant mutation (<Emphasis Type="Italic">Ms</Emphasis><Subscript><Emphasis Type="Italic">tc</Emphasis></Subscript>) derived from tissue culture

Male sterility mutations are an important tool in the investigation of anther and pollen development and for obtaining hybrid seeds in plant breeding. Cytological analysis of microsporo- and microgameto-genesis in sorghum plants with dominant mutation of male sterility (Ms_tc) derived from tissue culture has been carried out. Using substitution backcrosses, this mutation was introduced first into the nuclear background of the fertile sorghum line SK-723 and from this line into Volzhskoe-4w (V-4w). The mechanism of Ms_tc action on anther and pollen development differed in different nuclear backgrounds. In SK-723, phenotypic expression of Ms_tc began before the beginning of meiosis, which resulted in degeneration of sporogenous tissue in some anthers and in significant disturbances of anther morphology. In microsporocytes that did not degenerate, the frequency of non-specific meiotic abnormalities characteristic of the fertile line SK-723 significantly increased. In addition, in the mutant plants, a number of specific meiotic abnormalities--almost complete desynapsis, and formation of syncytial structures--were observed, apparently the consequence of Ms_tc action. In mono- or bi-nucleate microspores, degenerative processes resulting in formation of empty or anomalously coloured pollen grains led to almost complete male sterility. In the V-4w nuclear background, changes in anther structure and meiotic disturbances were infrequent. The degenerative processes began at the uni- or binucleate microspore stage and resulted in formation of empty or abnormally coloured pollen grains, and in partial pollen sterility. Thus, the same nuclear male sterility-inducing mutation in different nuclear backgrounds affects different stages of pollen development.     

Web-FACE: a new canopy free-air CO<Subscript>2</Subscript> enrichment system for tall trees in mature forests

The long-term responses of forests to atmospheric CO₂ enrichment have been difficult to determine experimentally given the large scale and complex structure of their canopy. We have developed a CO₂ exposure system that uses the free-air CO₂ enrichment (FACE) approach but was designed for tall canopy trees. The system consists of a CO₂-release system installed within the crown of adult trees using a 45-m tower crane, a CO₂ monitoring system and an automated regulation system. Pure CO₂ gas is released from a network of small tubes woven into the forest canopy (web-FACE), and CO₂ is emitted from small laser-punched holes. The set point CO₂ concentration ([CO₂]) of 500 µmol mol^-1 is controlled by a pulse-width modulation routine that adjusts the rate of CO₂ injection as a function of measured [CO₂] in the canopy. CO₂ consumption for the enrichment of 14 tall canopy trees was about 2 tons per day over the whole growing season. The seasonal daytime mean CO₂ concentration was 520 µmol mol^-1. One-minute averages of CO₂ measurements conducted at canopy height in the center of the CO₂-enriched zone were within ᆨ% and ᆞ% of the target concentration for 76% and 47% of the exposure time, respectively. Despite the size of the canopy and the windy site conditions, performance values correspond to about 75% of that reported for conventional forest FACE with the added advantage of a much simpler and less intrusive infrastructure. Stable carbon isotope signals captured by 80 Bermuda grass (Cynodon dactylon) seedlings distributed within the canopy of treated and control tree districts showed a clearly delineated area, with some nearby individuals having been exposed to a gradient of [CO₂], which is seen as added value. Time-integrated values of [CO₂] derived from the C isotope composition of C. dactylon leaves indicated a mean (-SD) concentration of 513ᇓ µmol mol^-1 in the web-FACE canopy area. In view of the size of the forest and the rough natural canopy, web-FACE is a most promising avenue towards natural forest experiments, which are greatly needed.     

Modeling the light interception and carbon gain of individual fluttering aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis> Michx) leaves

Poplars (Populus spp.) have a particular petiole morphology that enhances leaf flutter even in light winds. Previous studies have shown that this trait enhances whole canopy carbon gain through changes in the temporal dynamics and spatial distribution of light in the lower canopy. However, less is known about the effects of flutter for leaves at the top of the canopy ("sun leaves"). A computer simulation model was developed that uses latitude, time of day, day of year, azimuth and a slope component, which was varied at a 3 Hz frequency over an arc of rotation to create the flutter motion, and generate data on light interception for both surfaces of a fixed or fluttering leaf. The light files generated (10 Hz) were input into a dynamic model of photosynthesis to estimate the carbon gain for both fluttering and fixed leaves. As compared to leaves fixed at various angles and azimuths, fluttering leaves had a more uniform light interception. Depending on their angle and azimuth, fixed leaves may not always be intercepting high photon flux density (PFD) even when exposed to full sun. Leaf flutter continuously randomizes leaf angles creating uniform PFD inputs for photosynthetic reactions regardless of variation in leaf orientation and solar position. These effects on light interception could have positive impacts on carbon gain for leaves at the top of the canopy.