Carbon isotope discrimination in irrigated and droughted potato (Solanum tuberosum L.)

Carbon isotope discrimination (Δ) was measured in irrigated and droughted potato. Under irrigation, Δ in leaflets at given nodes increased (P < 0.001) between 21 and 63 d after emergence (DAE), which was attributed to increasing stomatal conductance (g_s) during leaf expansion. The effect of leaf position on Δ was non-significant in mature leaves. Under drought, Δ decreased (P < 0.001) in successive leaves up the stem, reflecting changes in g_s and water stress. At each node Δ remained constant or decreased, suggesting that effects of water stress were greater than changes with leaf expansion. There were significant differences in Δ between cultivars in both treatments, and in the progressive decrease in Δ up the stem under drought. Differences in Δ between cultivars were consistent with differences in stomatal control of leaf water status following water stress. Values for Δ in tubers were consistently lower than in stem and leaf, and decreased more rapidly. Differences in Δ between cultivars did not reflect dry matter production in either treatment, and differences in water use were non-significant between cultivars under drought. So, plants can achieve similar dry matter production through different growth strategies when irrigated or droughted, and Δ does not provide a simple, indirect method of selecting for dry matter production under water stress.     

Hydraulic conductance, stomatal conductance, and maximal photosynthetic rate in bean leaves

A positive correlation was found between steady state values of hydraulic (L _pA) and stomatal conductance (g _s) of French bean leaves: both were lower in the dark than in the light and lower in water-deficient plants than in the well-watered ones. The relative rate of stomatal opening after a pressure rise in the xylem was also positively related to L _pA. The L _pA and g _s were both related to the maximal photosynthetic rate at saturating CO2 concentrations. This revised version was published online in June 2006 with corrections to the Cover Date.     

Variations in photosynthetic rate and associated parameters with age of oil palm leaves under irrigation

Net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) in an adult oil palm (Elaeis guineensis) canopy were highest in the 9^th leaf and progressively declined with leaf age. Larger leaf area (LA) and leaf dry mass (LDM) were recorded in middle leaves. P _N showed a significant positive correlation with g _s and a negative relationship with leaf mass per area (ALM). The oil palm leaf remains photosynthetically active for a longer time in the canopy which contributes significantly to larger dry matter production in general and greater fresh fruit bunch yields in particular.     

Photosynthesis Inhibition During Gas Exchange Oscillations in ABA-Treated Helianthus annuus: Relative Role of Stomatal Patchiness and Leaf Carboxylation Capacity

Environmental factors that induce spatial heterogeneity of stomatal conductance, g _s, called stomatal patchiness, also reduce the photochemical capacity of CO₂ fixation, yet current methods cannot distinguish between the relative effect of stomatal patchiness and biochemical limitations on photosynthetic capacity. We evaluate effects of stomatal patchiness and the biochemical capacity of CO₂ fixation on the sensitivity of net photosynthetic rate (P _N) to stomatal conductance (g _s), θ (θ = δP _N/g _s). A qualitative model shows that stomatal patchiness increases the sensitivity θ while reduced biochemical capacity of CO₂ fixation lowers θ. We used this feature to distinguish between stomatal patchiness and mesophyll impairments in the photochemistry of CO₂ fixation. We compared gas exchange of sunflower (Helianthus annuus L.) plants grown in a growth chamber and fed abscisic acid, ABA (10⁻⁵ M), for 10 d with control plants (-ABA). P _N and g _s oscillated more frequently in ABA-treated than in control plants when the leaves were placed into the leaf chamber and exposed to a dry atmosphere. When compared with the initial CO₂ response measured at the beginning of the treatment (day zero), both ABA and control leaves showed reduced P _N at particular sub-stomatal CO₂ concentration (c _i) during the oscillations. A lower reduction of P _N at particular g _s indicated overestimation of c _i due to stomatal patchiness and/or omitted cuticular conductance, g _c. The initial period of damp oscillation was characterised by inhibition of chloroplast processes while stomatal patchiness prevailed at the steady state of gas exchange. The sensitivity θ remained at the original pre-treatment values at high g _s in both ABA and control plants. At low g _s, θ decreased in ABA-treated plants indicating an ABA-induced impairment of chloroplast processes. In control plants, g _c neglected in the calculation of g _s was the likely reason for apparent depression of photosynthesis at low g _s. This revised version was published online in August 2006 with corrections to the Cover Date.     

Coupled response of stomatal and mesophyll conductance to light enhances photosynthesis of shade leaves under sunflecks

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO₂ diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A_n), stomatal conductance (g_s) and mesophyll conductance (g_m) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower A_n, g_m, leaf nitrogen and photosynthetic capacity (A_max) but g_s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g_s and g_m increased, and A_n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of g_m with increased light enables shade leaves to respond quickly to sunflecks.     

Effects of root flooding and stage of development on the growth and photosynthesis of field bean (Vicia faba L. minor)

Field bean plants were subjected to flooding stress for 7 days, during two stages of development: at the vegetative phase (4-week-old seedlings) and at the generative phase (8-week-old plants). The height of plants, total area of leaves, the number of undamaged leaves, dry plant matter, chlorophyll content, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) activity, the maximum quantum yield of PS2 photochemistry (Fv/Fm ratio), the photosynthesis rate (P _N) and stomatal conductance (g _s) were determined. A strong reduction in stem elongation and leaf area as well as in dry matter production was observed as a result of flooding. The responses from vegetative plants were greater than in generative plants. Waterlogging decreased chlorophyll a and b in leaves, notably at the vegetative stage, and persisted after cessation of flooding. After flooding, photosynthesis was strongly reduced and positively correlated with decreased stomatal conductance. Damage to the photosynthetic apparatus resulted in a lower Fv/Fm especially in young seedlings. In vegetative plants Fv/Fm quickly returned to the control levels after the soil was drained. The results show that an excess of water in the soil limits growth and injures the photosynthetic apparatus in field beans, but that the extent of the injury is strongly age dependent.     

Over-expression of PaSOD in transgenic potato enhances photosynthetic performance under drought

Drought stress enhances the production of superoxide radical (O₂ ^._) and superoxide dismutase catalyses dismutation of it to H₂O₂ and O₂, and hence provides a first line of defense against oxidative stress. Over-expression of a cytosolic copper-zinc superoxide dismutase, cloned from Potentilla atrosanguinea (PaSOD), in potato (Solanum tuberosum ssp. tuberosum L. cv. Kufri Sutlej) resulted in enhanced net photosynthetic rates (P_N) and stomatal conductance (g_s) compared to that in the wild type (WT) plants under control (irrigated) as well as drought stress conditions. Drought stress declined leaf water potential, P_N, g_s, photosystem II activity, and chlorophyll content, but increased proline and O₂ ^._ content more in WT than transgenic potato plants (SS5). The significantly higher SOD activity in SS5 coincided well with lower O₂ ^._ content suggesting its role in maintaining higher g_s and P_N in transgenic potato plants.     

Disentangling the contributions of ontogeny and water stress to photosynthetic limitations in almond trees

Very few studies have attempted to disentangle the respective role of ontogeny and water stress on leaf photosynthetic attributes. The relative significance of both effects on photosynthetic attributes has been investigated in leaves of field‐grown almond trees [Prunus dulcis (Mill.) D. A. Webb] during four growth cycles. Leaf ontogeny resulted in enhanced leaf dry weight per unit area (W_a), greater leaf dry‐to‐fresh weight ratio and lower N content per unit of leaf dry weight (N_w). Concomitantly, area‐based maximum carboxylation rate (V_cmax), maximum electron transport rate (J_max), mesophyll conductance to CO₂ diffusion (gm)′ and light‐saturated net photosynthesis (A_max) declined in both well‐watered and water‐stressed almond leaves. Although g_m and stomatal conductance (g_s) seemed to be co‐ordinated, a much stronger coordination in response to ontogeny and prolonged water stress was observed between g_m and the leaf photosynthetic capacity. Under unrestricted water supply, the leaf age‐related decline of A_max was equally driven by diffusional and biochemical limitations. Under restricted soil water availability, A_max was mainly limited by g_s and, to a lesser extent, by photosynthetic capacity and g_m. When both ontogeny and water stress effects were combined, diffusional limitations was the main determinant of photosynthesis limitation, while stomatal and biochemical limitations contributed similarly.     

Hydraulic conductance,stomatal conductance,and maximal photosynthetic rate in bean leaves

A positive correlation was found between steady state values of hydraulic (L _pA) and stomatal conductance (g _s) of French bean leaves: both were lower in the dark than in the light and lower in water-deficient plants than in the well-watered ones. The relative rate of stomatal opening after a pressure rise in the xylem was also positively related to L _pA. The L _pA and g _s were both related to the maximal photosynthetic rate at saturating CO2 concentrations.     

Biomass partition,leaf gas exchange and water relations of alfalfa and milkvetch seedlings in response to soil drying

This study compared physiological and growth responses to water stress of two legume species during the seedling stage. Potted alfalfa (Medicago sativa L. cv. Algonquin) and milkvetch (Astragalus adsurgens Pall. cv. Pengyang earlymaturing vetch) seedlings were grown under well-watered [soil water content (SWC) maintained at 14.92% daily] or water-stressed conditions (drying) for 15 days. Net photosynthetic rate (P _N), transpiration rate (E) and stomatal conductance (g _s) of both species decreased parabolically. When SWC decreased to 7.2% and 10.3%, g _s values for alfalfa and milkvetch were significantly different from those of the respective well-watered plants (p<0.05). When SWC decreased to 6.6% for alfalfa and 6.8% for milkvetch, leaf water potentials (ψ_L) were significantly different from those of the well-watered plants (p<0.05). Thus the difference between the SWC thresholds for a nonhydraulic root signal (nHRS) and a hydraulic root signal (HRS) were 0.6% and 3.5% for alfalfa and milkvetch, respectively. Milkvetch had a lower g _s than alfalfa for a given SWC (p<0.05). Although alfalfa seedlings had a higher dry mass (DM) and root:shoot ratio (R/S) than milkvetch in both treatments (p<0.05), we concluded that milkvetch seedlings had greater drought tolerance than alfalfa.     

Interactions of Elevated CO2 Concentration and Drought Stress on Photosynthesis in Eucalyptus Cladocalyx F. Muell

Response of net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (c _i), and photosynthetic efficiency (F_v/F_m) of photosystem 2 (PS2) was assessed in Eucalyptus cladocalyx grown for long duration at 800 (C₈₀₀) or 380 (C₃₈₀) μmol mol^-1 CO₂ concentration under sufficient water supply or under water stress. The well-watered plants at C₈₀₀ showed a 2.2 fold enhancement of P _N without any change in g _s. Under both C₈₀₀ and C₃₈₀, water stress decreased P _N and g _s significantly without any substantial reduction of c _i, suggesting that both stomatal and non-stomatal factors regulated P _N. However, the photosynthetic efficiency of PS2 was not altered. This revised version was published online in June 2006 with corrections to the Cover Date.     

Stomatal behavior and photosynthetic performance under dynamic light regimes in a seasonally dry tropical rain forest

Rates of photosynthetic induction upon exposure to high light and rates of induction loss after darkening the leaf were measured in the field for four species of tropical shrubs in the family Rubiaceae. During wet season mornings, stomatal conductance (g _s) in the shade prior to induction was generally high enough so that the time course of induction was determined primarily by rates of activation of biochemical processes. During wet season afternoons, however, g _s values in the shade tended to be considerably lower and photosynthetic induction following a light increase exhibited a slower time course. In the afternoon, the time course of induction was determined by a combination of stomatal opening time and the rates of activation of light regulated enzymes. Stomatal behavior was also correlated with patterns of induction loss following a transfer from high light to darkness. In the afternoon, maximum g _s was lower for all species, and for a given time in the darkness, leaves showed a greater loss of induction in the afternoon than in the morning. During the dry season, maximum g _s and average values for g _s in the shade were reduced in all species. Along with these shifts in stomatal behavior, reduced rates of photosynthetic induction were observed. In the high-light species, the lower maximum g _s values observed during the dry season were also correlated with increased induction loss for a given time in the darkness. For all species, stomatal behavior was affected by season and time of day and, with the exception of wet season mornings, stomata appeared to exert significant control over rates of induction and patterns of induction loss. The results of simulation modeling suggest that the observed seasonal and diurnal changes in rates of induction and induction loss can have significant consequences on sunfleck carbon gain under a dynamic light regime. Received: 17 March 1999 / Accepted: 26 October 1999     

Determinant of photosynthetic capacity in rice leaves under ambient air conditions

At the grain-filling stage, net photosynthetic rate (P _N), stomatal conductance (g _s), and ribulose-1,5-bisphosphate carboxylation efficiency (CE) were correlated in order to find the determinant of photosynthetic capacity in rice leaves. For a flag leaf, P _N in leaf middle region was higher than in its upper region, and leaf basal region had the lowest P _N value. The differences in g _s and CE were similar. P _N, g _s, and CE gradually declined from upper to basal leaves, showing a leaf position gradient. The correlation coefficient between P _N and CE was much higher than that between P _N and g _s in both cases, and P _N was negatively correlated with intercellular CO₂ concentration (C _i). Hence the carboxylation activity or activated amount of ribulose-1,5-bisphosphate carboxylase/oxygenase rather than g_s was the determinant of the photosynthetic capacity in rice leaves. In addition, in flag leaves of different tillers P _N was positively correlated with g _s, but negatively correlated with C _i. Thus g _s is not the determinant of the photosynthetic capacity in rice leaves.The study was supported by the State Key Basic Research and Development Plan (No.G1998010100).     

Direct and acclimatory responses of stomatal conductance to elevated carbon dioxide in four herbaceous crop species in the field

In order to separate the net effect of growth at elevated [CO₂] on stomatal conductance (g_s) into direct and acclimatory responses, mid‐day values of g_s were measured for plants grown in field plots in open‐topped chambers at the current ambient [CO₂], which averaged 350 μmol mol^?1 in the daytime, and at ambient + 350 μmol mol^?1[CO₂] for winter wheat, winter barley, potato and sorghum. The acclimatory response was determined by comparing g_s measured at 700 μmol mol^?1[CO₂] for plants grown at the two [CO₂]. The direct effect of increasing [CO₂] from 350 to 700 μmol mol^?1 was determined for plants grown at the lower concentration. Photosynthetic rates were measured concurrently with g_s. For all species, growth at the higher [CO₂] significantly reduced g_s measured at 700 μmol mol^?1[CO₂]. The reduction in g_s caused by growth at the higher [CO₂] was larger for all species on days with low leaf to air water vapour pressure difference for a given temperature, which coincided with highest conductances and also the smallest direct effects of increased [CO₂] on conductance. For barley, there was no other evidence for stomatal acclimation, despite consistent down‐regulation of photosynthetic rate in plants grown at the higher [CO₂]. In wheat and potato, in addition to the vapour pressure difference interaction, the magnitude of stomatal acclimation varied directly in proportion to the magnitude of down‐regulation of photosynthetic rate through the season. In sorghum, g_s consistently exhibited acclimation, but there was no down‐regulation of photosynthetic rate. In none of the species except barley was the direct effect the larger component of the net reduction in g_s when averaged over measurement dates. The net effect of growth at elevated [CO₂] on mid‐day g_s resulted from unique combinations of direct and acclimatory responses in the various species.     

Carbon assimilation in contrasting streamside and inland Spartina alterniflora salt marsh

Carbon assimilation and standing crop biomass of Spartina alterniflora were studied in a contrasting streamside and inland salt marsh in Louisiana Gulf coast, USA. A substantially lower leaf dry weight, leaf area index, and standing crop biomass were recorded for inland plants as compared to streamside plants. Net assimilation rates ranged between 8 to 25 mol m^–2 s^–1 for streamside and between 4 to 19 mol m^–2 s^–1 for inland plants. The average photosynthetic rates were significantly lower for inland plants which were growing in an apparently more stressed environment. In addition, the differences were more profound with progression of the growing season. The reduced photosynthetic activity in the inland marsh was attributed to greater soil waterlogging, increased anaerobic root respiration, plant toxins (sulfide), restricted nutrient uptake or a combination of these factors.Abbreviations E_h = redox potential - g_w = stomatal conductance - LAI = leaf area index - P_n = net photosynthesis - PPFD = photosynthetic photon flux density - T₁ = leaf temperature     

Water-use efficiency of a mallee eucalypt growing naturally and in short-rotation coppice cultivation

This study compared mature Eucalyptus kochii subsp. plenissima trees in inner regions or edges of natural bushland to young trees belt-planted through cleared agricultural land as uncut saplings or regenerating coppice over 2.7 years at Kalannie, Western Australia (320 mm annual rainfall). We assessed the ability of the species to alter its gas exchange characteristics, leaf physical attributes, and water-use efficiency of foliar carbon assimilation (WUE _i) or of total dry matter production (WUE _DM). Stomatal conductance (g _s) varied five-fold between treatment means, with coppices exhibiting greatest values and mature bush least. Photosynthetic rates followed this trend. Leaf photosynthetic capacity estimated by chlorophyll content varied 1.3-fold parallel with variations in leaf thickness, with coppices rating lowest and mature edge trees most highly. WUE _i varied 1.5-fold between treatments and was greatest in mature inner-bush and edge trees. Leaf photosynthetic capacity and g _s were both correlated with WUE _i. Carbon isotope composition (δ¹³C values) of new shoot dry matter produced early in a seasonal flush were similar to those of root starch but when averaged over the whole season correlated well with WUE _i and gas exchange characteristics of trees of each treatment. Coppices showed poorest WUE _i and most negative shoot tip δ¹³C but their WUE _DM was high. This discrepancy was suggested to relate to carbon allocation strategies in coppices favouring fast growth of replacement shoots but not of roots. Physiology of coppice growth of E. kochii is usefully geared towards both rapid and water-use efficient production of woody biomass in water limited environments.     

Photosynthetic and growth responses of <Emphasis Type="Italic">Camelina sativa</Emphasis> (L.) Crantz to varying nitrogen and soil water status

Water and nitrogen (N) deficiency are two major constraints limiting the yield and quality of many oilseed crops worldwide. This study was designed to assess the response of Camelina sativa (L.) Crantz to the availability of N and water resources on photosynthesis and yield parameters. All the measured variables, which included plant height, root and shoot dry matter, root:shoot ratio, xylem pressure potential (XPP), yield components, photosynthetic parameters, and instantaneous water-use efficiency (WUE) were remarkably influenced by water and nitrogen supply. Net photosynthetic rate (P _N) and yield components were significantly decreased more by water deficit than by N deficiency. XPP, stomatal conductance (g _s), and intercellular CO₂ concentration (C _i) decreased substantially as the water deficit increased irrespective of the level of N application. WUE at the high N supply [100 and 150 kg(N) ha⁻¹] dropped in a large degree as the increased water deficit due to a larger decrease in P _N than transpiration rate (E). The results of this study suggest that the regulative capacity of N supply on photosynthetic and plant growth response is significantly affected by soil water status and C. sativa is more sensitive to water deficit than N supply.     

Photosynthetic characteristics of dwarf and fringe Rhizophora mangle L. in a Belizean mangrove

Twin Cays (Belize) is a highly oligotrophic mangrove archipelago dominated by Rhizophora mangle L. Ocean‐fringing trees are 3–7 m tall with a leaf area index (LAI) of 2.3, whereas in the interior, dwarf zone, trees are 1.5 m or less, and the LAI is 0.7. P‐fertilization of dwarf trees dramatically increases growth. As a partial explanation of these characteristics, it was hypothesized that differences in stature and growth rates would reflect differences in leaf photosynthetic capacity, as determined by the photochemical and biochemical characteristics at the chloroplast level. Gas exchange and chlorophyll fluorescence were used to compare photosynthesis of dwarf, fringe and fertilized trees. Regardless of zonation or treatment, net CO₂ exchange (A) and photosynthetic electron transport were light saturated at less than 500 µmol photons m^?2 s^?1, and low‐light quantum efficiencies were typical for healthy C₃ plants. On the other hand, light‐saturated A was linearly related to stomatal conductance (g_s), with seasonal, zonal and treatment differences in photosynthesis corresponding linearly to differences in the mean g_s. Overall, photosynthetic capacity appeared to be co‐regulated with stomatal conductance, minimizing the variability of C_i at ambient CO₂ (and hence, C_i/C_a). Based on the results of in vitro assays, regulation of photosynthesis in R. mangle appeared to be accomplished, at least in part, by regulation of Rubisco activity.     

Effects of elevated CO2 on leaf gas exchange in beech and oak at two levels of nutrient supply: consequences for sensitivity to drought in beech

Beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) were grown from seed for two whole seasons at two CO₂ concentrations (ambient and ambient + 250 μmol mol^?1) with two levels of soil nutrient supply. Measurements of net leaf photosynthetic rate (A) and stomatal conductance (g_s) of well-watered plants were taken over both seasons; a drought treatment was applied in the middle of the second growing season to a separate sample of beech drawn from the same population. The net leaf photosynthetic rate of well-watered plants was stimulated in elevated CO₂ by an average of 75% in beech and 33% in oak; the effect continued through both growing seasons at both nutrient levels. There were no interactive effects of CO₂ concentration and nutrient level on A or g_s in beech or oak. Stomatal conductance was reduced in elevated CO₂ by an average of 34% in oak, but in beech there were no significant reductions in g_s except under cloudy conditions (–22% in elevated CO₂). During drought, there was no effect of CO₂ concentration on g_s in beech grown with high nutrients, but for beech grown with low nutrients, g_s was significantly higher in elevated CO₂, causing more rapid soil drying. With high nutrient supply, soil drying was more rapid at elevated CO₂ due to increased leaf area. It appears that beech may substantially increase whole-plant water consumption in elevated CO₂, especially under conditions of high temperature and irradiance when damage due to high evaporative demand is most likely to occur, thereby putting itself at risk during periods of drought.     

Photosynthetic characteristics of dipterocarp species planted on degraded sandy soils in southern Thailand

To elucidate whether dipterocarp species, dominant late-successional species of tropical forests in Southeast Asia, actually have a disadvantage when planted on open site in terms of their photosynthetic characteristics, we investigated photosynthesis in dipterocarp seedlings planted in the open on degraded sandy soils in southern Thailand. These species were compared with seedlings of Acacia mangium Willd., a fast-growing tropical leguminous tree, which is often planted on degraded open site in Southeast Asia. The dipterocarp seedlings had an irradiance-saturated net photosynthetic rate (P _N), stomatal conductance (g _s), carboxylation efficiency, and photosynthetic capacity comparable to or superior to those of A. mangium. In particular, seedlings of Dipterocarpus obtusifolius Teijsm. ex Miq. showed an irradian-ce-saturated P _N of 21 μmol m⁻² s⁻¹, a value higher than any previously reported for a dipterocarp species, accompanied by high g _s (0.7 mol m⁻² s⁻¹) and high photosynthetic capacity. Thus dipterocarp species do not necessarily have a disadvantage in terms of their photosynthetic characteristics on open sites with degraded sandy soils.