Stomatal conductance and not stomatal density determines the long-term reduction in leaf transpiration of poplar in elevated CO<Subscript>2</Subscript>

Using a free-air CO₂ enrichment (FACE) experiment, poplar trees (Populus × euramericana clone I214) were exposed to either ambient or elevated [CO₂] from planting, for a 5-year period during canopy development, closure, coppice and re-growth. In each year, measurements were taken of stomatal density (SD, number mm⁻²) and stomatal index (SI, the proportion of epidermal cells forming stomata). In year 5, measurements were also taken of leaf stomatal conductance (g _s, μmol m⁻² s⁻¹), photosynthetic CO₂ fixation (A, mmol m⁻² s⁻¹), instantaneous water-use efficiency (A/E) and the ratio of intercellular to atmospheric CO₂ (C_i:C_a). Elevated [CO₂] caused reductions in SI in the first year, and in SD in the first 2 years, when the canopy was largely open. In following years, when the canopy had closed, elevated [CO₂] had no detectable effects on stomatal numbers or index. In contrast, even after 5 years of exposure to elevated [CO₂], g _s was reduced, A/E was stimulated, and C_i:C_a was reduced relative to ambient [CO₂]. These outcomes from the long-term realistic field conditions of this forest FACE experiment suggest that stomatal numbers (SD and SI) had no role in determining the improved instantaneous leaf-level efficiency of water use under elevated [CO₂]. We propose that altered cuticular development during canopy closure may partially explain the changing response of stomata to elevated [CO₂], although the mechanism for this remains obscure.     

Contrasting leaf characteristics of trees and lianas in secondary and mature forests in southwestern China

We compared variation in sun-canopy leaf anatomy, morphology and photosynthetic rates of coexisting woody species (trees and lianas) in an 8-year-old secondary forest (SF) and mature forest (MF) in the wet season in Xishuangbanna, SW China. Variability of leaf traits of 66 species within growth-form groups in each forest was quantified using coefficients of variation (CV). For the mean values, the woody species in the SF had significantly higher leaf thickness and stomatal density, but lower nonmesophyll/mesophyll ratios than those in the MF. The average leaf area and leaf mass area (LMA) in the studied woody species did not change greatly during the successional process, but differed significantly between the growth forms, with trees having higher values than lianas. The light-saturated photosynthetic rate per unit leaf area (A _a) of the woody species in the SF ranged from 11.2 to 34.5 μmol m⁻² s⁻¹, similarly to pioneer tree species from literature data in southeast Asia. The A _a and photosynthetic nitrogen-use efficiency (PNUE) were significantly higher than those in the MF; whereas A _a in the MF ranged between 9 to 21 μmol m⁻² s⁻¹, with similar values between lianas and trees. For all woody species in both SF and MF, there were no significant differences in the average values of the CV of all measured variables for both lianas and trees. However, considerable variation in leaf anatomy, morphology, and photosynthetic rates within both growth forms and forests existed, as well as substantial variation in leaf size and stomatal density. We concluded that the tropical woody species formed a heterogeneous functional group in terms of leaf morphology and physiology in both secondary and mature forests.     

<Emphasis Type="Italic">Petunia</Emphasis> × <Emphasis Type="Italic">hybrida</Emphasis> during transition to flowering as affected by light intensity and quality treatments

Petunia × hybrida was grown under high (H), medium (M) and low (L) light intensity [photoperiod; 16 h d⁻¹, photosynthetic photon flux density (PPFD); 360, 120 and 40 μmol m⁻² s⁻¹, respectively] as well as under end-of-day (EOD) red (R) and far-red (FR) light quality treatments [photoperiod; 14.5 h d⁻¹, PPFD; 30 μmol m⁻² s⁻¹ EOD; 15 min, Control (C) light; without EOD light treatment]. Shoot growth, leaf anatomical and photosynthetic responses as well as the responses of peroxidase (POD) isoforms and their specific activities following transition to flowering (1–6 weeks) were evaluated. Flower bud formation of Petunia × hybrida was achieved at the end of the 4th week for H light treatment and on the end of the 6th week for FR light treatment. No flower bud formation was noticed in the C and R light treatments. H and M light treatments induced lower chlorophyll (Chla, Chlb, Chla+b) concentrations in comparison to L light. On the other hand R and FR light chlorophyll content were similar to C light. Photosynthetic parameters [CO₂ assimilation rate (A), transpiration rate (E) and stomatal conductance (g _s) values] were higher in the H light treated plants in comparison to M and L light treated plants. A, E and g _s values of R and FR light were similar to C light plants. Leaf anatomy revealed that total leaf thickness, thickness of the contained tissues (epidermis, palisade and spongy parenchyma) and relative volume percentages of the leaf histological components were differently affected within the light intensity and the light quality treatments. POD specific activities increased from the 1st to the 6th week during transition to flowering. Native-PAGE analysis revealed the appearance of four anionic POD (A₁–A₄) isoforms in all light treatments. On the basis of the leaf anatomical, photosynthetic and plant morphological responses, the production of high quality Petunia × hybrida plants with optimal flowering times could be achieved through the control of both light intensity and light quality. The appearance of A₁ and A₂ anionic POD isoforms could be also used for successful scheduling under light treatments.     

Photosynthetic and leaf anatomical characteristics of Castanea sativa: a comparison between in vitro and nursery plants

The anatomic and functional leaf characteristics related to photosynthetic performance of Castanea sativa growing in vitro and in nursery were compared. The irradiance saturated photosynthesis in in vitro grown plantlets was significantly lower compared to nursery plants (65 vs. 722 μmol m⁻² s⁻¹). The maximum photosynthetic rate (P_Nmax) was 4.0 and 10.0 μmol(CO₂) m⁻² s⁻¹ in in vitro microshoots and nursery plant leaves, respectively. Carboxylation efficiency (C_E) and electron transport rate (ETR) were three-folds higher in nursery plants than in microshoots. The nonphotochemical quenching (NPQ) was saturated at 80 μmol m⁻² s⁻¹ in microshoots suggesting limited photoprotection by thermal dissipation. The microshoots had wide open, spherical stomata and higher stomatal density than nursery plants and they had almost no epicuticular wax. Consequently, the microshoots had high stomatal conductance and high transpiration rate. These anatomic and functional leaf characteristics are likely major causes of the low survival rates of plantlets after ex vitro transfer.     

Response of photosynthesis to high light and drought for Arabidopsis thaliana grown under a UV-B enhanced light regime

Arabidopsis thaliana grown in a light regime that included ultraviolet-B (UV-B) radiation (6 kJ m⁻² d⁻¹) had similar light-saturated photosynthetic rates but up to 50% lower stomatal conductance rates, as compared to plants grown without UV-B radiation. Growth responses of Arabidopsis to UV-B radiation included lower leaf area (25%) and biomass (10%) and higher UV-B absorbing compounds (30%) and chlorophyll content (52%). Lower stomatal conductance rates for plants grown with UV-B radiation were, in part, due to lower stomatal density on the adaxial surface. Plants grown with UV-B radiation had more capacity to down regulate photochemical efficiency of photosystem II (PSII) as shown by up to 25% lower φ_PSII and 30% higher non-photochemical quenching of chlorophyll fluorescence under saturating light. These contributed to a smaller reduction in the maximum photochemical efficiency of PSII (F _v/F _m), greater dark-recovery of F _v/F _m, and higher light-saturated carbon assimilation and stomatal conductance and transpiration rates after a four-hour high light treatment for plants grown with UV-B radiation. Plants grown with UV-B were more tolerant to a 12 day drought treatment than plants grown without UV-B as indicated by two times higher photosynthetic rates and 12% higher relative water content. UV-B-grown plants also had three times higher proline content. Higher tolerance to drought stress for Arabidopsis plants grown under UV-B radiation may be attributed to both increased proline content and decreased stomatal conductance. Growth of Arabidopsis in a UV-B-enhanced light regime increased tolerance to high light exposure and drought stress.     

Genotypic variation in photosynthesis in cacao is correlated with stomatal conductance and leaf nitrogen

Variation in photosynthetic parameters was observed between eight contrasting cacao (Theobroma cacao) genotypes. Net photosynthetic rate (P_N) ranged from 3.4 to 5.7 μmol(CO₂) m⁻² s⁻¹ for the genotypes IMC 47 and SCA 6, respectively. Furthermore, genotypic differences were detected in quantum efficiency ranging from 0.020 to 0.043 μmol(CO₂) μmol⁻¹(photon) for UF 676 and AMAZ 15/15, respectively. Differences in PN were correlated with both stomatal conductance (g_s) and leaf nitrogen per unit area. Some variation in water use efficiency was observed between genotypes, both intrinsic (P_N/g_s) and instantaneous (P_N/transpiration rate). Both measures of water use efficiency were a negative function of specific leaf area. Evidence was found for a trade-off mechanism between cacao genotypes in photosynthesis and leaf structure. High photosynthetic rate, expressed on a mass basis was associated with smaller leaves. Furthermore, thinner leaves were compensated for by a higher nitrogen content per unit mass.     

Long-time variations in leaf mass and area of Mediterranean evergreen broad-leaf and narrow-leaf maquis species

Morphological (dry mass, DM; surface area, LA; leaf mass per area, LMA), anatomical (leaf thickness, L), phenological (leaf life span, LL), and physiological (net photosynthetic rate, P _N) leaf traits of the evergreen species co-occurring in the Mediterranean maquis developing at Castelporziano (Rome) were tested. The correlation analysis indicated that LMA variation was tightly associated with LL variations: Cistus incanus and Arbutus unedo had a short LL (4±1, summer leaves, and 11±1 months, respectively) and low LMA (153±19 g m⁻²) values, Quercus ilex, Phillyrea latifolia, and Pistacia lentiscus high LMA (204±7 g m⁻²) and long LL (22±3 months), Erica arborea, Erica multiflora, and Rosmarinus officinalis a short LL (9±2 months) and an either high (213±29 g m⁻², R. officinalis and E. multiflora) or low (115±17 g m⁻², E. arborea) LMA. LMA values were significantly (p≤0.05) correlated with P _N (r≥0.68). In the tested species, LMA increased in response to the decrease of the total rainfall during the leaf expansion period. LMA variation was due to the unequal variation of DM and LA in the considered species. LMA is thus a good indicator of evergreen maquis species capability to respond to climate change, in particular to total rainfall decrease in the Mediterranean basin.     

Effects of soil flooding and changes in light intensity on photosynthesis of <Emphasis Type="Italic">Eugenia uniflora</Emphasis> L. seedlings

The increased frequency of heavy rains as a result of global climate change can lead to flooding and changes in light availability caused by the presence of thick clouds. To test the hypothesis that reduction in light availability can alleviate the harmful effects of soil flooding on photosynthesis, the authors studied the effects of soil flooding and acclimation from high to low light on the photosynthetic performance of Eugenia uniflora. Seedlings acclimated to full sunlight (about 35 mol m⁻² d⁻¹) for 5 months were transferred to partial sunlight (about 10 mol m⁻² d⁻¹) and were either subjected to soil flooding or not flooded. Chlorophyll fluorescence was measured throughout the flooding period and leaf gas exchange was measured 16 days after flooding was initiated. Minimal fluorescence yield (Fo) was significantly higher and the quantum efficiency of open PSII centres (Fv/Fm) was significantly lower in flooded than in non-flooded plants in full sunlight. Sixteen days after flooding was initiated, stomatal conductance (gs_sat) and net photosyntheses expressed on a leaf area (A_sat-area), weight (A_sat-wt) and chlorophyll (A_sat-Chl) basis decreased in response to soil flooding. Flooding decreased stomatal conductance by similar amounts in full and partial sunlight, but A_sat-area in partial and full sunlight was 3.4 and 16.8 times lower, respectively, in flooded than in non-flooded plants. These results indicate that changes from full to partial sunlight during soil flooding can alleviate the effects of flooding stress on photosynthesis in E. uniflora seedlings acclimated to full sunlight. The responses of photosynthesis in trees to flooding stress may be dependent on changes in light environment during heavy rains.     

Leaf chlorosis in oilseed rape plants (Brassica napus) grown on cadmium-polluted soil: causes and consequences for photosynthesis and growth

Brassica napus L. (oilseed rape) was grown from seeds on a reconstituted soil contaminated with cadmium (100 mg Cd kg⁻¹ dry soil), resulting in a marked chlorosis of the leaves which was investigated using a combination of biochemical, biophysical and physiological methods. Spectroscopic and chromatographic analyses of the photosynthetic pigments indicated that chlorosis was not due to a direct interaction of Cd with the chlorophyll biosynthesis pathway. In addition, mineral deficiency and oxidative stress were apparently not involved in the pigment loss. Leaf chlorosis was attributable to a marked decrease in the chloroplast density caused by a reduction in the number of chloroplasts per cell and a change in cell size, suggesting that Cd interfered with chloroplast replication and cell division. Relatively little Cd was found in the chloroplasts and the properties of the photosynthetic apparatus (electron transport, protein composition, chlorophyll antenna size, chloroplast ultrastructure) were not affected appreciably in plants grown on Cd-polluted soil. Depth profiling of photosynthetic pigments by phase-resolved photoacoustic spectroscopy revealed that the Cd-induced decrease in pigment content was very pronounced at the leaf surface (stomatal guard cells) compared to the leaf interior (mesophyll). This observation was consistent with light transmission and fluorescence microscopy analyses, which revealed that stomata density in the epidermis was noticeably reduced in Cd-exposed leaves. Concomitantly, the stomatal conductance estimated from gas-exchange measurements was strongly reduced with Cd. When plants were grown in a high-CO₂ atmosphere (4,000 μl CO₂ l⁻¹), the inhibitory effect of Cd on growth was not cancelled, suggesting that the reduced availability of CO₂ at the chloroplast level associated with the low stomatal conductance was not the main component of Cd toxicity in oilseed rape. Received: 14 July 2000 / Accepted: 27 August 2000     

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     

Is distribution of hydraulic constraints within tree crowns reflected in photosynthetic water-use efficiency? An example of <Emphasis Type="Italic">Betula pendula</Emphasis>

Spatial and daily variation in photosynthetic water-use efficiency was examined in leaves of Betula pendula Roth with respect to distribution of hydraulic conductance within the crown, morphological properties of stomata, and water availability. Intrinsic water-use efficiency (A _n/g _s) was determined from gas-exchange measurements performed both in situ in a natural forest stand and on detached shoots under laboratory conditions. In intact foliage, sun leaves demonstrated significantly higher (P < 0.001) A _n/g _s than shade leaves, as photosynthesis in the lower canopy was chronically limited by low light availability. However, this difference reversed in the mid-day period under sufficient irradiance (I > 800 μmol m⁻² s⁻¹): A _n/g _s averaged 28.8 and 24.0 μmol mol⁻¹ (P < 0.01) for shade and sun leaves, respectively. This last finding coincided with the data obtained in laboratory conditions: under equivalent leaf water supply and light, A _n/g _s in shade foliage was greater (P < 0.001) than in sun foliage across a wide range of irradiance. Thus, shade foliage of B. pendula is characterized by inherently higher A _n/g _s than sun foliage, associated with more conservative stomatal behavior, and lower soil-to-leaf (K _T) and leaf hydraulic conductances. Under unlimited light conditions, a within-crown trade-off between A _n/g _s and K _T becomes apparent. Differences in stomatal conductance between the detached shoots from sunlit and shaded canopy layers were largely attributable to the variation in stomatal morphology; significant relationships were established with characteristics combining stomatal size and density (relative stomatal surface, stomatal pore area index). Stomatal morphology is very likely involved in long-term adjustment of photosynthetic WUE.     

Photosynthetic responses of Microstegium vimineum (Trin.) A. Camus, a shade-tolerant, C4 grass, to variable light environments

Microstegium vimineum (Trin.) A. Camus, a shade-tolerant C₄ grass, has spread throughout the eastern United States since its introduction in 1919. This species invades disturbed understory habitats along streambanks and surrounding mesic forests, and has become a major pest in areas such as Great Smoky Mountains National Park. The focus of this study was to characterize the photosynthetic induction responses of M. vimineum, specifically its ability to utilize low light and sunflecks, two factors that may be critical to invasive abilities and survival in the understory. In addition, we were curious about the ability of a grass with the C₄ photosynthetic pathway to respond to sunflecks. Plants were grown under 25% and 50% ambient sunlight, and photosynthetic responses to both steady-state and variable light were determined. Plants grown in both 25% and 50% ambient sun became 90% light saturated between 750–850 μmol m⁻² s⁻¹; however, plants grown in 50% ambient sun had significantly higher maximum steady-state photosynthetic rates (16.09 ± 1.37 μmol m⁻² s⁻¹ vs. 12.71 ± 1.18 μmol m⁻² s⁻¹). Both groups of plants induced to 50% of the steady-state rate in 3–5 min, while it took 10–13 min to reach 90% of maximum rates, under both flashing and steady-state light. For both groups of plants, stomatal conductance during induction reached maximum rates in 6–7 min, after which rates decreased slightly. Upon return to low light, rates of induction loss and stomatal closure were very rapid in both groups of plants, but were more rapid in those grown in high light. Rapid induction and the ability to induce under flashing light may enable this species to invade and dominate mesic understory habitats, while rapid induction loss due to stomatal closure may prevent excess water loss when low light constrains photosynthesis. The C₄ pathway itself does not appear to present an insurmountable barrier to the ability of this grass species to respond to sunflecks in an understory environment. Received: 21 February 1997 / Accepted: 10 October 1997     

Sensitivity of Metasequoia glyptostroboides to ozone stress

2-year-old seedlings of Metasequoia glyptostroboides were grown in open top chambers and exposed to four ozone concentrations [O₃] (charcoal-filtered air, CF; 50, 100, and 200 mm³ m⁻³) for 25 d. Measurements of growth, leaf chlorophyll (Chl) content, and gas exchange parameters were made before and/or after O₃ exposure. Leaf length, crown width, Chl a/b, net photosynthetic rate, stomatal conductance, and transpiration rate were significantly reduced at 100 and 200 mm³(O₃) m⁻³. A remarkable decrease in stomatal conductance also occurred at 50 mm³(O₃) m⁻³.     

Chlorophyll fluorescence imaging of photosynthetic activity in sun and shade leaves of trees

The differences in pigment levels, photosynthetic activity and the chlorophyll fluorescence decrease ratio R _Fd (as indicator of photosynthetic rates) of green sun and shade leaves of three broadleaf trees (Platanus acerifolia Willd., Populus alba L., Tilia cordata Mill.) were compared. Sun leaves were characterized by higher levels of total chlorophylls a + b and total carotenoids x + c as well as higher values for the weight ratio chlorophyll (Chl) a/b (sun leaves 3.23–3.45; shade leaves: 2.74–2.81), and lower values for the ratio chlorophylls to carotenoids (a + b)/(x + c) (with 4.44–4.70 in sun leaves and 5.04–5.72 in shade leaves). Sun leaves exhibited higher photosynthetic rates P _N on a leaf area basis (mean of 9.1–10.1 μmol CO₂ m⁻² s⁻¹) and Chl basis, which correlated well with the higher values of stomatal conductance G _s (range 105–180 mmol m⁻² s⁻¹), as compared to shade leaves (G _s range 25–77 mmol m⁻² s⁻¹; P _N: 3.2–3.7 μmol CO₂ m⁻² s⁻¹). The higher photosynthetic rates could also be detected via imaging the Chl fluorescence decrease ratio R _Fd, which possessed higher values in sun leaves (2.8–3.0) as compared to shade leaves (1.4–1.8). In addition, via R _Fd images it was shown that the photosynthetic activity of the leaves of all trees exhibits a large heterogeneity across the leaf area, and in general to a higher extent in sun leaves than in shade leaves.     

Soil salinity effects on transpiration and net photosynthetic rates,stomatal conductance and Na+ and Cl− contents in durum wheat

Leaf gas exchange, plant growth and leaf ion content were measured in wheat (Triticum durum L. cv. HD 4502) exposed to steady- state salinities (1.6, 12.0 and 16.0 dS nr⁻¹) for 8 weeks. Salinity reduced leaf area and number of tillers, and increased Na⁺ and Cl⁻ concentrations in leaves. Leaf- to- leaf gradients of these ions were observed. The oldest leaf contained 6 to 8 times more Na⁺ and Cl⁻ than the flag leaf. Net photosynthetic rate (P_N), transpiration rate (E) and stomatal conductance (g_S) were the highest in flag leaf, declined in the middle and fully expanded leaves, and were minimum in the oldest leaves. These processes were reduced by salinity with similar leaf- to- leaf gradients. Intercellular CO₂ concentrations in the older leaves were higher than in the flag leaf in non-saline plants, and increased similarly with salinity. Leaf age was the major factor in reducing P_N, and senescence processes were promoted by salinity.     

Gas-exchange analysis of chloroplastic fructose-1,6-bisphosphatase antisense potatoes at different air humidities and at elevated CO2

Gas-exchange measurements were performed to analyze the leaf conductances and assimilation rates of potato (Solanum tuberosum L. cv. Desireé) plants expressing an antisense construct against chloroplastic fructose-1,6-bisphosphatase (FBPase, EC 3.1.3.11) in response to increasing photon flux densities, different relative air humidities and elevated CO₂ concentrations. Assimilation rates (A) and transpiration rates (E) were observed during a stepwise increase of photon flux density. These experiments were carried out under atmospheric conditions and in air containing 500 μmol mol⁻¹ CO₂. In both gas atmospheres, two levels of relative air humidity (60–70% and 70–80%) were applied in different sets of measurements. Intercellular CO₂ concentration, leaf conductance, air-to-leaf vapour pressure deficit, and instantaneous water-use efficiency (A/E) were determined. As expected, assimilation rates of the FBPase antisense plants were significantly reduced as compared to the wild type. Saturation of assimilation rates in transgenic plants occurred at a photon flux density of 200 μmol m⁻² s⁻¹, whereas saturation in wild type plants was observed at 600 μmol m⁻² s⁻¹. Elevated ambient CO₂ levels did not effect assimilation rates of transgenic plants. At 70–80% relative humidity and atmospheric CO₂ concentration the FBPase antisense plants had significantly higher leaf conductances than wild-type plants while no difference emerged at 60–70%. These differences in leaf conductance vanished at elevated levels of ambient CO₂. Stomatal response to different relative air humidities was not affected by mesophyll photosynthetic activity. It is suggested that the regulation of stomatal opening upon changes in photon flux density is merely mediated by a signal transmitted from mesophyll cells, whereas the intercellular CO₂ concentration plays a minor role in this kind of stomatal response. The results are discussed with respect to stomatal control by environmental parameters and mesophyll photosynthesis. Received: 24 September 1998 / Accepted: 9 February 1999     

Unusually low carbon isotope ratios in plants from hanging gardens in southern Utah

Leaf carbon isotope ratios (δ¹³C) and photosynthetic gas exchange were measured on plants growing in hanging garden communities in southern Utah, USA. Hanging gardens are unusual, mesic cliff communities occurring where water seeps from the sandstone bedrock in an otherwise extremely arid region; there is very limited overlap in species distributions inside and outside these gardens. Solar exposure in hanging gardens varied with orientation and one of the gardens (Ribbon Garden) was shaded throughout the day. The leaf δ¹³C values of plants in hanging gardens were significantly more negative than for plants from either nearby ephemeral wash or riparian communities. In Ribbon Garden, the observed δ¹³C values were as low as −34.8‰, placing them among the most negative values reported for any terrestrial plant species growing in a natural environment. Hanging garden plants were exposed to normal atmospheric CO₂ with an average δ¹³C value of −7.9‰ and so the low leaf δ¹³C values could not be attributed to exposure to a CO₂ source with low ¹³C content. There was a seasonal change toward more negative leaf δ¹³C values at the end of the growing season. The observed leaf δ¹³C values were consistent with photosynthetic gas exchange measurements that indicated unusually high leaf intercellular CO₂ concentrations associated with the relatively low light levels in hanging gardens. Thus, extremely negative leaf δ¹³C values would be expected if significant amounts of the seasonal carbon gain occur at light levels low enough to be near the light compensation point. Maximum observed photosynthetic rates varied with light levels at each of the gardens, with maximum rates averaging 20.3, 14.6, and 3.1 μmol m⁻²s⁻¹ at Double Garden, Lost Garden, and Ribbon Garden, respectively. Leaf nitrogen contents averaged 18.5 mg g⁻¹ in species from the more shaded hanging gardens (Lost and Ribbon). When expressed on a leaf area basis, nitrogen contents averaged 117 mmol N m⁻² at Lost Garden and 65 mmol N m⁻² at Ribbon Garden (shadiest of the two gardens). Leaf nitrogen isotope ratios averaged −2.3‰ (range of −0.7 to −6.1‰), suggesting that most of the nitrogen was derived from a biological fixation source which is most likely the Nostoc growing on the sandstone walls at the seep. These values contrast with leaf nitrogen isotope ratios of 5–9‰ which have been previously reported for arid zone plants in nearby ecosystems. Received: 19 January 1997 / Accepted: 19 April 1997     

Alterations in Rubisco activity and in stomatal behavior induce a daily rhythm in photosynthesis of aerial leaves in the amphibious-plant Nuphar lutea

Nuphar lutea is an amphibious plant with submerged and aerial foliage, which raises the question how do both leaf types perform photosynthetically in two different environments. We found that the aerial leaves function like terrestrial sun-leaves in that their photosynthetic capability was high and saturated under high irradiance (ca. 1,500 μmol photons m⁻² s⁻¹). We show that stomatal opening and Rubisco activity in these leaves co-limited photosynthesis at saturating irradiance fluctuating in a daily rhythm. In the morning, sunlight stimulated stomatal opening, Rubisco synthesis, and the neutralization of a night-accumulated Rubisco inhibitor. Consequently, the light-saturated quantum efficiency and rate of photosynthesis increased 10-fold by midday. During the afternoon, gradual closure of the stomata and a decrease in Rubisco content reduced the light-saturated photosynthetic rate. However, at limited irradiance, stomatal behavior and Rubisco content had only a marginal effect on the photosynthetic rate, which did not change during the day. In contrast to the aerial leaves, the photosynthesis rate of the submerged leaves, adapted to a shaded environment, was saturated under lower irradiance. The light-saturated quantum efficiency of these leaves was much lower and did not change during the day. Due to their low photosynthetic affinity for CO₂ (35 μM) and inability to utilize other inorganic carbon species, their photosynthetic rate at air-equilibrated water was CO₂-limited. These results reveal differences in the photosynthetic performance of the two types of Nuphar leaves and unravel how photosynthetic daily rhythm in the aerial leaves is controlled.     

The photosynthetic characteristics of differently shaped leaves in Populus euphratica Olivier

There is a distinct leaf shape polymorphism within a single plant of P. euphratica Olivier. The anatomical structure, carbon isotope discrimination (Δ¹³C), and stomatal and photosynthetic behaviour were investigated in broad-ovate (BOL) and lanceolate (LL) leaves, located at the top and bottom in crown, respectively, of a mature Euphrates poplar growing in its native habitat. Both types of leaves had a non-Kranz anatomy and low Δ¹³C values. However, Δ¹³C of a LL was in average 3.2‰ larger than that of a BOL. In comparison with the LL, the BOL had a smaller stomatal conductance, causing subsequent decreases in transpiration rate and ratio of CO₂ concentrations in intercellular spaces to air. Carbon assimilation rate and water use efficiency were higher in the BOLs than in the LLs. The BOL exhibited C₄-like enzymological features, the activity of glycollate oxidase, and the ratio of activities of ribulose-1,5-bisphosphate carboxylase (RuBPC) to phosphoenolpyruvate carboxylase (PEPC) was lower in BOL than in LL throughout the whole growing season. The lowered ratio of RuBPC/PEPC in BOL was mainly associated with a marked decline in the activity of RuBPC, and only a slight increase in the activity of PEPC. These differences might contribute to microclimate adaptation in both types of leaves. This revised version was published online in June 2006 with corrections to the Cover Date.     

In vitro growth and single-leaf photosynthetic response of Cymbidium plantlets to super-elevated CO2 under cold cathode fluorescent lamps

To examine the effectiveness of super-elevated (10,000 μmol mol⁻¹) CO₂ enrichment under cold cathode fluorescent lamps (CCFL) for the clonal propagation of Cymbidium, plantlets were cultured on modified Vacin and Went (VW) medium under 0, 3,000 and 10,000 μmol mol⁻¹ CO₂ enrichment and two levels of photosynthetic photon flux density (PPFD, 45 and 75 μmol m⁻² s⁻¹). Under high PPFD, 10,000 μmol mol⁻¹ CO₂ increased root dry weight and promoted shoot growth. In addition, a decrease in photosynthetic capacity and chlorosis at leaf tips were observed. Rubisco activity and stomatal conductance of these plantlets were lower than those of plantlets at 3,000 μmol mol⁻¹ CO₂ under high PPFD, which had a higher photosynthetic capacity. On the other hand, plantlets on Kyoto medium grown in 10,000 μmol mol⁻¹ CO₂ under high PPFD had a higher photosynthetic rate than those on modified VW medium; no chlorosis was observed. Furthermore, growth of plantlets, in particular the roots, was remarkably enhanced. This result indicates that a negative response to super-elevated CO₂ under high PPFD could be improved by altering medium components. Super-elevated CO₂ enrichment of in vitro-cultured Cymbidium could positively affect the efficiency and quality of commercial production of clonal orchid plantlets.