Woody tissue photosynthesis and its contribution to trunk growth and bud development in young plants

Stem photosynthesis can contribute significantly to woody plant carbon balance, particularly in times when leaves are absent or in ‘open’ crowns with sufficient light penetration. We explored the significance of woody tissue (stem) photosynthesis for the carbon income in three California native plant species via measurements of chlorophyll concentrations, radial stem growth, bud biomass and stable carbon isotope composition of sugars in different plant organs. Young plants of Prunus ilicifolia, Umbellularia californica and Arctostaphylos manzanita were measured and subjected to manipulations at two levels: trunk light exclusion (100 and 50%) and complete defoliation. We found that long‐term light exclusion resulted in a reduction in chlorophyll concentration and radial growth, demonstrating that trunk assimilates contributed to trunk carbon income. In addition, bud biomass was lower in covered plants compared to uncovered plants. Excluding 100% of the ambient light from trunks on defoliated plants led to an enrichment in ¹³C of trunk phloem sugars. We attributed this effect to a reduction in photosynthetic carbon isotope discrimination against ¹³C that in turn resulted in an enrichment in ¹³C of bud sugars. Taken together our results reveal that stem photosynthesis contributes to the total carbon income of all species including the buds in defoliated plants.     

Seasonal dynamics of non-structural carbohydrates in two species of mediterranean sub-shrubs with different leaf phenology

The seasonal dynamics of non-structural carbohydrates in the woody organs of two co-existing mediterranean sub-shrubs were analyzed. The two species show different leaf phenology during summer: Linum suffruticosum, maintains many of its green leaves, while Lepidium subulatum sheds most of its leaves. These different leaf phenologies are related to different strategies with regard to summer stress. The maintenance of leaves in Linum is related to its stress tolerance while Lepidium avoids stress by shedding its leaves. The main objectives were to: (1) determine the differences in the seasonal dynamics of non-structural carbohydrates among the main woody organs of both species; (2) verify if differences in the leaf phenology, and hence in the strategy with regard to summer drought, lead to different seasonal patterns of carbohydrate storage and use between the two species; (3) compare the seasonal dynamics of carbohydrates of the two studied sub-shrubs with those of mediterranean trees and shrubs previously reported in the literature. The concentration of soluble sugars (SS), starch and total non-structural carbohydrates (TNC) were assessed monthly, over 17 months, in the main roots, stems and the transition zone between root and shoot systems of both species. Starch storage capacity and SS, starch and TNC pools were calculated. The seasonal pattern of carbohydrate accumulation was similar among the woody organs analyzed, but it differed with those reported for mediterranean trees and shrubs. The two species showed different pools and seasonal patterns of non-structural carbohydrate concentrations in its woody organ, which corresponded to their different extent of leaf shedding. The stress-avoider Lepidium accumulated starch during spring shoot growth as a carbon store for summer respiration and had low pools of SS, whereas the stress-tolerant Linum increased SS during summer drought to maintain photosynthetic activity during summer and had low starch pools and storage capacity. However, irrespective of their different leaf shedding patterns, both species had a similar relative variation of their TNC concentration, which contrasts with previous results on deciduous and evergreen woody species.     

Shifts in bryophyte carbon isotope ratio across an elevation?��?soil age matrix on Mauna Loa, Hawaii: do bryophytes behave like vascular plants?

The carbon isotope ratio (δ(13)C) of vascular plant leaf tissue is determined by isotope discrimination, primarily mediated by stomatal and mesophyll diffusion resistances and by photosynthetic rate. These effects lead to predictable trends in leaf δ(13)C across natural gradients of elevation, irradiance and nutrient supply. Less is known about shifts in δ(13)C for bryophytes at landscape scale, as bryophytes lack stomata in the dominant gametophyte phase, and thus lack active control over CO(2) diffusion. Twelve bryophyte species were sampled across a matrix of elevation and soil ages on Mauna Loa, Hawaii Island. We tested hypotheses based on previous findings for vascular plants, which tend to have less negative δ(13)C at higher elevations or irradiances, and for leaves with higher leaf mass per area (LMA). Across the matrix, bryophytes spanned the range of δ(13)C values typical of C(3) vascular plants. Bryophytes were remarkably similar to vascular plants in exhibiting less negative δ(13)C with increasing elevation, and with lower overstory cover; additionally δ(13)C was related to bryophyte canopy projected mass per area, a trait analogous to LMA in vascular plants, also correlated negatively with overstory cover. The similarity of responses of δ(13)C in bryophytes and vascular plants to environmental factors, despite differing morphologies and diffusion pathways, points to a strong direct role of photosynthetic rate in determining δ(13)C variation at the landscape scale.     

Fractionation of Carbon Isotopes during Biogenesis of Atmospheric Isoprene

The stable carbon isotope composition of isoprene emitted from leaves of red oak (Quercus rubra L.) was measured. Isoprene was depleted in ¹³C relative to carbon recently fixed by photosynthesis. The difference in isotope composition between recently fixed carbon and emitted isoprene was independent of the isotopic composition of the source CO₂. β-Carotene, an isoprenoid plant constituent, was depleted in ¹³C relative to whole leaf carbon to the same degree as isoprene, but fatty acids were more depleted. Isoprene emitted from leaves fed abscisic acid was much less depleted in ¹³C than was isoprene emitted from unstressed leaves. We conclude that isoprene is made from an isoprenoid precursor that is derived from acetyl-CoA made from recent photosynthate. The carbon isotope composition of isoprene in the atmosphere is likely to be slightly more negative (less ¹³C) than C₃ plant material but when plants are stressed the isotopic composition could vary.     

Carbon isotope composition as a tool to control the quality of herbs and medicinal plants

Isotope screening is a simple test for determining the photosynthetic pathway used by plants. The scope of this work was to classify the photosynthetic type of some herbs and medicinal plants through studies of the carbon isotope composition (δ¹³C). Also, we propose the use of carbon isotope composition as a tool to control the quality of herbs and medicinal plants. For studies of δ¹³C, δ¹³C‰ = [R (sample)/R (standard) − 1] × 10⁻³, dry leaves powdered in cryogenic mill were analyzed in a mass spectrometer coupled with an elemental analyzer for determining the ratio R = ¹³CO₂/¹²CO₂. In investigation of δ¹³C of 55 species, 23 botanical families, and 44 species possessed a C₃ photosynthetic type. Six species found among the botanical families Euphorbiaceae and Poaceae were C₄ plants, and 5 species found among the botanical families Agavaceae, Euphorbiaceae, and Liliaceae possessed CAM-type photosynthesis. Carbon isotope composition of plants can be used as quality control of herbs and medicinal plants, allowing the identification of frauds or contaminations. Also, the information about the photosynthetic type found for these plants can help in introducing and cultivating exotic and wild herbs and medicinal plants.     

亚热带季风常绿阔叶林植被恢复Ⅱ．──木本豆科植物的气体交换

测定了几种木本豆科植物和乡土树种如木荷、桂西荷的叶片光合速率月变化,并在７和１２月测定了几种乡土树种、木本豆科植物和亚热带常绿阔叶树的气体交换量．结果表明,乡土树种在５月或９月光合速率达最大值,而在５—９月之间由于太阳辐射强烈和气温较高,光合速率降低;但木本豆科植物绢毛相思和大叶相思光台速率则较高,说明木本豆科植物在华南夏季能适应当地酷热条件,保持较高光合速率．在７月,阴香日同化量为８．５９９·ｍ－２,马占相思为５．６９９·ｍ－２,乡土树种木荷达７．４６９·ｍ－２．由此可见,乡土树种或常绿阔叶树种与木本豆科植物的阔叶混交林有助于提高太阳辐射能利用和增加光合同化产物,从而有利于亚热带常绿阔叶林植被的恢复．     

Photosynthetic and gas exchange characteristics of dominant woody plants on a moisture gradient in an African savanna

We determined key photosynthetic gas exchange parameters, and their temperature dependence, in dominant woody plants at four savanna sites on a moisture gradient in Botswana, southern Africa. Leaf stable carbon and nitrogen (N) isotope and morphological measures were made concurrently. Sampling of these predominantly non‐N‐fixing species took place during an exceptional rainfall season, representing near‐optimum conditions for primary production at these sites. The mean specific leaf area and leaf size were positively related to mean annual rainfall (MAR); species with larger leaves of lower density were more abundant in wetter sites. Almost all species at all sites showed high net light‐saturated photosynthetic rates (A_max?10 μmol CO₂ m^?2 s^?1) due both to high CO₂ carboxylation (V_c,max) and RubP‐regeneration capacity (J_max). These high rates were associated with high values of leaf [N]. Across all sites, the temperature response of A_max showed no clear optimum, and a gradual drop from 25°C to 35°C, without notable temperature limitation at leaf temperatures in excess of 35°C. Dark respiration rate (R_day) across all species and sites increased exponentially with increasing leaf temperature. Species sampled at selected sites revealed a negative relationship between leaf δ¹³C (stable carbon isotope ratio) and MAR, suggesting higher leaf‐level water‐use efficiency at drier sites when integrated over the life of the leaf. At wetter sites, specific leaf [N] was lower and photosynthetic nitrogen‐use efficiency increased, a pattern reflected at the ecosystem level by less ¹⁵N enrichment of leaves at these sites. Taken together, the results suggest a switch from water‐use to nitrogen‐use efficiency constraints with increasing moisture availability. These constraints impact leaf form and function significantly, and may emerge at the ecosystem level in aspects of water and N cycling.     

Nutritional regulation in mixotrophic plants: new insights from Limodorum abortivum

Partially mycoheterotrophic (mixotrophic) plants gain carbon from both photosynthesis and their mycorrhizal fungi. This is considered an ancestral state in the evolution of full mycoheterotrophy, but little is known about this nutrition, and especially about the physiological balance between photosynthesis and fungal C gain. To investigate possible compensation between photosynthesis and mycoheterotrophy in the Mediterranean mixotrophic orchid Limodorum abortivum, fungal colonization was experimentally reduced in situ by fungicide treatment. We measured photosynthetic pigments of leaves, stems, and ovaries, as well as the stable C isotope compositions (a proxy for photosynthetic C gain) of seeds and the sizes of ovaries and seeds. We demonstrate that (1) in natural conditions, photosynthetic pigments are most concentrated in ovaries; (2) pigments and photosynthetic C increase in ovaries when fungal C supply is impaired, buffering C limitations and allowing the same development of ovaries and seeds as in natural conditions; and (3) responses to light of pigment and ¹³C contents in ovaries shift from null responses in natural conditions to responses typical of autotrophic plants in treated L. abortivum, demonstrating photoadaptation and enhanced use of light in the latter. L. abortivum thus preferentially feeds on fungi in natural conditions, but employs compensatory photosynthesis to buffer fungal C limitations and allow seed development.     

Carbonic anhydrase and its influence on carbon isotope discrimination during C4 photosynthesis. Insights from antisense RNA in Flaveria bidentis

In C4 plants, carbonic anhydrase (CA) facilitates both the chemical and isotopic equilibration of atmospheric CO2 and bicarbonate (HCO3-) in the mesophyll cytoplasm. The CA-catalyzed reaction is essential for C4 photosynthesis, and the model of carbon isotope discrimination (Delta13C) in C4 plants predicts that changes in CA activity will influence Delta13C. However, experimentally, the influence of CA on Delta13C has not been demonstrated in C4 plants. Here, we compared measurements of Delta13C during C4 photosynthesis in Flaveria bidentis wild-type plants with F. bidentis plants with reduced levels of CA due to the expression of antisense constructs targeted to a putative mesophyll cytosolic CA. Plants with reduced CA activity had greater Delta13C, which was also evident in the leaf dry matter carbon isotope composition (delta13C). Contrary to the isotope measurements, photosynthetic rates were not affected until CA activity was less than 20% of wild type. Measurements of Delta13C, delta13C of leaf dry matter, and rates of net CO2 assimilation were all dramatically altered when CA activity was less than 5% of wild type. CA activity in wild-type F. bidentis is sufficient to maintain net CO2 assimilation; however, reducing leaf CA activity has a relatively large influence on Delta13C, often without changes in net CO2 assimilation. Our data indicate that the extent of CA activity in C4 leaves needs to be taken into account when using Delta13C and/or delta13C to model the response of C4 photosynthesis to changing environmental conditions.     

Scaling sun and shade photosynthetic acclimation of Alocasia macrorrhiza to whole-plant performance – I. Carbon balance and allocation at different daily photon flux densities

We determined the carbon allocation patterns and construction costs of Alocasia macrorrhiza plants grown at different photon flux densities (PFD) as well as the whole-plant carbon gain of these plants at different daily PFDs. Growth at high PFD resulted in thicker leaves with a higher leaf mass per unit area, and increased biomass allocation to petioles and roots, as compared to growth at low PFD. Increased allocation to petioles may have been necessary to support the heavier leaves, whereas increased allocation to roots may have been necessary to supply sufficient water for the higher transpiration rates in high PFD. Root biomass was highly correlated with the daily, whole-plant transpiration rate. Tissue construction costs per unit dry mass were unchanged by acclimation, but, since the mass per unit areas of leaves, roots and petioles all increased, construction costs per unit leaf area were much higher for plants grown at high PFD. On a per unit leaf area basis, daily whole-plant carbon gain measured at high daily PFD was higher in high- than in low-PFD-grown plants. However, on a per unit leaf mass basis, low-PFD-grown plants had a daily carbon gain at least as high as that of high-PFD-grown plants at high daily PFD. At low daily PFD, low-PFD-grown plants maintained an advantage over high-PFD-grown plants in terms of carbon gain because of their larger leaf area ratios. Thus, in terms of carbon gain, low-PFD-grown plants performed better than sun plants at low PFD and as well as high-PFD-grown plants at high PFD, despite their lower photosynthetic capacities per unit area. For high-PFD-grown plants, the higher construction costs per unit leaf area resulted in lower leaf area ratios, which counteracted the advantage of higher photosynthetic rates per unit leaf area.     

Natural abundance of 15N and 13C in nodulated legumes and other plants in the cerrado and neighbouring regions of Brazil

Leaves from over 1000 Brazilian native plants growing in the cerrado and neighbouring regions were sampled for C and N content. Half of these were analysed for ¹⁵N and further samples for ¹³C and ash content. Nodulated legumes from all three sub-families were included, together with two types of reference plant, non-nodulated legumes and non-legumes. Particular emphasis was placed on the large caesalpinioid genus Chamaecrista which is here for the first time reported to fix nitrogen in its native habitats. Woody and herbaceous species of this and other nodulated genera, with the exception of the mimosoid tree Stryphnodendron, showed evidence of nitrogen fixation. Amounts fixed were site-specific as was the ¹⁵N signature of reference plants. There was no evidence that nodulated legumes had higher leaf N than non-nodulated legumes: both were higher than non-legumes. Several species of Chamaecrista from section absus and species of Stryphnodendron had carbon contents of 50–55%, higher than previously reported for leaves. This was coupled with low (1–3%) ash contents. The ¹³C values of plants with 49% C were significantly more negative than those with <49% C: most species in the former group were woody and most in the latter group herbaceous. Mimosa pudica was unusual in having a wide range of percent C, percent ash and ¹³C values; these parameters were significantly correlated. It is concluded that Brazilian native legumes can fix significant amounts of nitrogen in the nutrient-poor cerrado soils. Consideration of mineral and lipid nutrition will be necessary in order fully to understand relations between ¹³C, carbon content and other physiological parameters.     

Temporal variation of δ<Superscript>13</Superscript>C of larch leaves from a montane boreal forest in Mongolia

This paper reports the temporal variation (2002–2004) in foliar δ¹³C values, which are indicative of long-term integrated photosynthetic and water use characteristics, of Siberian larch (Larix sibirica Ledeb.) trees in a montane forest at Mongonmorit, NE Mongolia. At the stand, the δ¹³C value for understory shaded leaves was more negative by 2‰ on average than that for sunlit leaves sampled concurrently from open and sun-exposed environments in a forest gap. The δ¹³C value of both sunlit and shaded leaves showed pronounced intra- but relatively small inter-seasonal variations. The δ¹³C value was more positive for juvenile than mature leaves. We conjecture that juvenile leaves may derive carbon reserves in woody tissues (e.g., stems). Regardless of leaf habitats, the δ¹³C value was also affected by insect herbivores occurred in mid summer of 2003, being more negative in newly emerging leaves from the twigs after defoliation than in non-defoliated mature leaves. This pattern seems to contrast with that for the juvenile leaves in the early growing season. We surmise that the newly emerging leaves used stored organic carbon that was depleted due to fractionation during remobilization and translocation for leaf regrowth. There was also intra- and inter-seasonal variation in the foliar N concentrations and C:N ratios. A good positive (negative) correlation between the foliar δ¹³C values and N concentrations (C:N ratios) was also observed for both sunlit and shaded leaves, suggesting that the relationship between water and nitrogen use is a crucial factor affecting the plant carbon–water relationship in this mid latitude forest with a cold semiarid climate. Our isotopic data demonstrate that the larches in NE Mongolia exhibits relatively higher water use efficiency with a distinct within-season variability.     

Covariation in leaf and root traits for native and non-native grasses along an altitudinal gradient in New Zealand

Across 30 grassland sites in New Zealand that ranged from native alpine grasslands to low elevation improved pastures, there were consistent patterns of leaf and root traits and significant differences between native and non-native grasses. Plants of high altitude sites have low N concentrations in both their leaves and roots, have thick leaves and roots, yet no differences in tissue density or photosynthetic water use efficiency when compared to plants of low altitude sites. Both the leaves and roots of the low altitude plants were enriched in (15)N relative to the plants of higher altitude, indicating that the low-N set of traits is associated with a more closed N cycle at high altitude. A second independent set of correlations shows that plants of wetter habitats have lower photosynthetic water use efficiency (more negative partial differential (13)C) and lower leaf and root tissue density than the plants of drier sites. For both leaves and roots, plants of native species consistently had traits associated with lower resource availability: lower N concentrations, denser tissues, more negative partial differential (15)N, and more positive partial differential (13)C than non-native species. If root %N is correlated with root longevity as has been shown in other systems, root longevity may be able to be predicted from simple measurements of leaf %N, though a hysteresis in the relationship between leaf and root N concentrations may make prediction of high longevity roots difficult.     

Does leaf shedding increase the whole-plant carbon gain despite some nitrogen being lost with shedding?

When old leaves are shed, part of the nitrogen in the leaf is retranslocated to new leaves. This retranslocation will increase the whole-plant carbon gain when daily C gain : leaf N ratio (daily photosynthetic N-use efficiency, NUE) in the old leaf, expressed as a fraction of NUE in the new leaf, becomes lower than the fraction of leaf N that is resorbed before shedding (R(N)). We examined whether plants shed their leaves to increase the whole-plant C gain in accord with this criterion in a dense stand of an annual herb, Xanthium canadense, grown under high (HN) and low (LN) nitrogen availability. The NUE of a leaf at shedding expressed as a fraction of NUE in a new leaf was nearly equal to the R(N) in the LN stand, but significantly lower than the R(N) in the HN stand. Thus shedding of old leaves occurred as expected in the LN stand, whereas in the HN stand, shedding occurred later than expected. Sensitivity analyses showed that the decline in NUE of a leaf resulted primarily from a reduction in irradiance in the HN stand. On the other hand, it resulted from a reduction in irradiance and also in light-saturated photosynthesis : leaf N content ratio (potential photosynthetic NUE) in the LN stand.     

Use of decreasing foliar carbon isotope discrimination during water limitation as a carbon tracer to study whole plant carbon allocation

Foliar carbon isotope discrimination (Δ) of C₃ plants decreases in water‐deficit situations as discrimination by the photosynthetic primary carboxylation reaction decreases. This diminished Δ in leaves under water deficit can be used as a tracer to study whole plant carbon allocation patterns. Carbon isotope composition (δ¹³C value) of leaf hot water extracts or leaf tissue sap represents a short‐term integral of leaf carbon isotope discrimination and thus represents the δ¹³C value of source carbon that may be distributed within a plant in water‐deficit situations. By plotting the δ¹³C values of source carbon against the δ¹³C values of sink tissues, such as roots or stems, it is possible to assess carbon allocation to and incorporation into sink organs in relation to already present biomass. This natural abundance labelling method has been tested in three independent experiments, a one‐year field study with the fruit tree species Ziziphus mauritiana and peach (Prunus persica), a medium‐term drought stress experiment with Ziziphus rotundifolia trees in the glasshouse, and a short‐term drought stress experiment with soybean (Glycine max). The data show that the natural abundance labelling method can be applied to qualitatively assess carbon allocation in drought‐stressed plants. Although it is not possible to estimate exact fluxes of assimilated carbon during water deficit the method represents an easy to use tool to study integrated plant adaptations to drought stress. In addition, it is a less laborious method that can be applied in field studies as well as in controlled experiments, with plants from any developmental stage.     

Carbon isotope ratios and the variation in the diurnal pattern of malate accumulation in aerial roots of CAM species of <Emphasis Type="Italic">Phalaenopsis</Emphasis> (Orchidaceae)

We investigated the carbon isotope ratios and the diurnal pattern of malate accumulation in leaves and aerial roots of eight species of Phalaenopsis grown in greenhouses. The leaves of all the species showed carbon isotope ratios and the diurnal patterns of malate content typical of CAM plants. However, the aerial roots exhibited a large variation in the diurnal pattern of malate content among species and even among plants within the same species, although carbon isotope ratios were always CAM-like values. Some aerial roots showed the typical diurnal pattern of CAM, but others maintained high or low malate contents during a day without fluctuation. In order to characterize more strictly the nature of the malate variation in the aerial roots, we further investigated a possible variation of the diurnal pattern of malate among different aerial roots within an individual for Phalaenopsis amabilis and P. cornu-cervi. The diurnal pattern of malate content was varied even among different aerial roots within the same plant. Thus the photosynthetic carbon metabolism in aerial roots of orchids is fairly complex.     

Variable carbon isotope ratios of Dudleya species growing in natural environments

Summary Measurements are reported of carbon isotope ratios of Dudleya species growing in natural plant communities. Considerable variation in the ¹³C values are interpreted as indicating substantial flexibility of the photosynthetic pathways between C₃ and CAM. The variability in photosynthetic pathway was in response to genetic factors, stage of plant development, life-form, and environmental conditions. Species active during drought periods have less carbon isotope fractionation than species that are summer-dormant. Summer-active species from drier habitats have less negative ¹³C values than those from more mesic sites. On the same plant, leaf tissue had more negative ¹³C values than tissue from the inflorescence. The less negative carbon isotope ratios are indicative of an increased proportion of exogenous CO₂ fixed in dark vs light. The ecological significance of these results is discussed.CIW-DPB Publication No. 522     

Foliar δ13C Within Temperate Deciduous Forest: Its Spatial Change and Interspecies Variation

The stable carbon isotope ratios (δ13C) of dominant tree and shrub species in typical temperate broad-leaved forests in Dongling Mountain, northwestern part of Beijing, China, were studied. The results indicated that the values of foliar 8 13C differed among the species tested, in the following order of Prunus armeniaca var. ansu Maxim. > Fraxinus rhychophylla Hce. ≈ Virex negundo var. heterophylla (Franch.) ReM. ≈ Quercus liaotungensis Koiz. ≈ Syringa pekinensis Rupr. > Juglans mandshurica Maxim. The δ13C values were also affected by the maturity of leaves. The newly-developed leaves had higher δ13C values than the mature ones, especially in Juglans mandshurica and Quercus liaotungensis. The conditions of habitats, particularly soil depth and moisture content, also had pronounced effect on foliar δ13C. The foliar δ13C values of plants grown in dry habitats were higher than those in moist ones. They were also affected by the position of leaves in the canopy: leaves at upper canopy showed higher δ13C values than the lower ones. It is suggested that stable isotope technique may be effective for the study of ecophysiology of plants in the warm temperate regions of China.     

δ(13) C of leaf-respired CO(2) reflects intrinsic water-use efficiency in barley

Leaf intrinsic water-use efficiency (WUE), the ratio of photosynthetic rate to stomatal conductance (A/g(s) ), is a key plant trait linking terrestrial carbon and water cycles. A rapid, integrative proxy for A/g(s) is of benefit to crop breeding programmes aiming to improve WUE, but also for ecologists interested in plant carbon-water balance in natural systems. We hypothesize that the carbon isotope composition of leaf-respired CO(2) (δ(13) C(Rl) ), two hours after leaves are transferred to the dark, records photosynthetic carbon isotope discrimination and so provides a proxy for A/g(s) . To test this hypothesis, δ(13) C(Rl) was measured in four barley cultivars grown in the field at two levels of water availability and compared to leaf-level gas exchange (the ratio of leaf intercellular to ambient CO(2) partial pressure, C(i) /C(a) , and A/g(s) ). Leaf-respired CO(2) was more (13) C-depleted in plants grown at higher water availability, varied between days as environmental conditions changed, and was significantly different between cultivars. A strong relationship between δ(13) C(Rl) and δ(13) C of sucrose was observed. δ(13) C(Rl) was converted into apparent photosynthetic discrimination (Δ(13) C(Rl) ) revealing strong relationships between Δ(13) C(Rl) and C(i) /C(a) and A/g(s) during the vegetative stage of growth. We therefore conclude that δ(13) C(Rl) may provide a rapid, integrative proxy for A/g(s) in barley.