Enhance Carbon Gain Research Articles

CAM-physiology and carbon gain of the orchid Phalaenopsis in response to light intensity, light integral and CO2.

The regulation of photosynthesis and carbon gain of crassulacean acid metabolism (CAM) plants has not yet been disclosed to the extent of C3-plants. In this study, the tropical epiphyte Phalaenopsis cv. "Sacramento" was subjected to different lighting regimes. Photosynthesis and biochemical measuring techniques were used to address four specific questions: (1) the response of malate decarboxylation to light intensity, (2) the malate carboxylation pathway in phase IV, (3) the response of diel carbon gain to the light integral and (4) the response of diel carbon gain to CO2 . The four CAM-phases were clearly discernable. The length of phase III and the malate decarboxylation rate responded directly to light intensity. In phase IV, CO2 was initially mainly carboxylated via Rubisco. However, at daylength of 16 h, specifically beyond ±12 h, it was mainly phosphoenolpyruvate carboxylase (PEP-C) carboxylating CO2 . Diel carbon gain appeared to be controlled by the light integral during phase III rather than the total daily light integral. Elevated CO2 further enhanced carbon gain both in phase IV and phase I. This establishes that neither malate storage capacity, nor availability of PEP as substrate for nocturnal CO2 carboxylation were limiting factors for carbon gain enhancement. These results advance our understanding of CAM-plants and are also of practical importance for growers.

Cytogenetic analysis of Bienertia sinuspersici Akhani as the first step in genome sequencing.

C4 plants are efficient in suppressing photorespiration and enhancing carbon gain as compared to C3 plants. Bienertia sinuspersici Akhani is one of the few species in the family Amaranthaceae that can perform C4 photosynthesis within individual chlorenchyma cells, without the conventional Kranz anatomy in its leaf. This plant is salt-tolerant and is well-adapted to thrive in hot and humid climates. To date, there have been no reported cytogenetic analyses yet on this species. This study aims to provide a cytogenetic analysis of B. sinuspersici as the first step in genome sequencing. Fluorescence in situ hybridization (FISH) karyotype analysis was conducted using the metaphase chromosomes of B. sinuspersici probed with 5S and 45S rDNA and Arabidopsis-type telomeric repeats. Results of the cytogenetic analysis confirmed that B. sinuspersici has 2n = 2x = 18 consisting of nine pairs of metacentric chromosomes. Two loci of 45S rDNA were found on the distal regions of the short arm of chromosome 7. Nine loci of 5S rDNA were found in the pericentromeric regions of chromosomes 1, 3, 4, 6, and 8, which also colocalized with Arabidopsis-type telomeric repeats; while four loci in the interstitial regions of chromosome 5 and 8 can be observed. The single locus of 5S rDNA that was found in chromosome 8 appears to be hemizygous. The FISH karyotype analysis, based on the combination of rDNAs, telomeric tandem repeat markers and C0t DNA chromosome landmarks, allowed efficient chromosome identification and provided useful information in characterizing the genome of B. sinuspersici.

Genetic dissection of photochemical efficiency under water-deficit stress in rice

Chlorophyll fluorescence (Chl-F) measurements together with non-invasive estimations of chlorophyll content, can be used to investigate functionally rich or poor photosystem II (PSII), relating to alterations in photosynthetic performances under different abiotic stresses. The aim was to identify genetic loci that control rice capacity to cope with different soil moisture conditions such as non-stress (control), water deficit and recovery during the reproductive stage. A genome-wide association study was performed for effective quantum yield of photosystem II (QY) and chlorophyll index across all three treatments. Accessions showed significant variability in traits within each treatment. A total of 43 genetic loci associated with QY and chlorophyll index were identified. Of the total genetic loci identified, 14 were for control, 13 for water-deficit stress and 16 for recovery responses. Interestingly, the majority of the identified genetic loci were co-localized either with chlorophyll synthesis or degradation pathways, components of PSII, transcription factors, protein kinases, transporters, kinases, and antioxidants genes. Favorable alleles and donor accessions found in our study would complement efforts aimed at stacking of traits. Moreover, our results provide promising genetic information for future validation and a potential resource for improving photochemical efficiency and subsequently enhancing carbon gain in rice under water-limited conditions.

Comparative Chloroplast Genome Analysis of Single-Cell C 4 Bienertia Sinuspersici with Other Amaranthaceae Genomes

Bienertia sinuspersici is a single-cell C4 (SCC4) plant species whose photosynthetic mechanisms occur in two cytoplasmic compartments containing central and peripheral chloroplasts. The efficiency of the C4 photosynthetic pathway to suppress photorespiration and enhance carbon gain has led to a growing interest in its research. A comparative analysis of B. sinuspersici chloroplast genome with other genomes of Amaranthaceae was conducted. Results from a 70% cut off sequence identity showed that B. sinuspersici is closely related to Beta vulgaris with slight variations in the arrangement of few genes such ycf1 and ycf15; and, the absence of psbB in Beta vulgaris. B. sinuspersici has the largest 153, 472 bp while Spinacea oleracea has the largest protein-coding sequence 6,754 bp larger than B. sinuspersici. The GC contents of each of the species ranges from 36.3 to 36.9% with B. sinuspersici having the same GC percentage as Haloxylon persicum and H. ammodendron (36.6%). The IR size also varies yet in all six species, the Ira/LSC border is generally located upstream of the trnH-GUG gene. A total of 107 tandem repeats were found in each of the species, most of which are situated in the intergenic space. These results provide basic information that may be valuable for future related studies.

Nitrogen-controlled intra- and interspecific competition between Populus purdomii and Salix rehderiana drive primary succession in the Gongga Mountain glacier retreat area.

In this study, intra- and interspecific competition were investigated in early successional Salix rehderiana Schneider and later-appearing Populus purdomii Rehder under non-fertilized (control) and nitrogen (N)-fertilized conditions in the Hailuogou glacier retreat area. Our aim was to discover whether N is a key factor in plant-plant competition and whether N drives the primary succession process in a glacier retreat area. We analyzed differences in responses to intra- and interspecific competition and N fertilization between P. purdomii and S. rehderiana, including parameters such as biomass accumulation, nutrient absorption, non-structural carbohydrates, photosynthetic capacity, hydrolysable amino acids and leaf ultrastructure. In the control treatments, S. rehderiana individuals subjected to interspecific competition benefited from the presence of P. purdomii plants, as indicated by higher levels of biomass accumulation, photosynthetic capacity, N absorption, amino acid contents and photosynthetic N-use efficiency. However, in the N-fertilized treatments, P. purdomii individuals exposed to interspecific competition benefited from the presence of S. rehderiana plants, as shown by a higher growth rate, enhanced carbon gain capacity, greater amino acid contents, and elevated water-use efficiency, whereas the growth of S. rehderiana was significantly reduced. Our results demonstrate that N plays a pivotal role in determining the asymmetric competition pattern among Salicaceae species during primary succession. We argue that the interactive effects of plant-plant competition and N availability are key mechanisms that drive primary succession in the Gongga Mountain glacier retreat area.

Impaired Stomatal Control Is Associated with Reduced Photosynthetic Physiology in Crop Species Grown at Elevated [CO2

Physiological control of stomatal conductance (Gs) permits plants to balance CO2-uptake for photosynthesis (PN) against water-loss, so optimizing water use efficiency (WUE). An increase in the atmospheric concentration of carbon dioxide ([CO2]) will result in a stimulation of PN and reduction of Gs in many plants, enhancing carbon gain while reducing water-loss. It has also been hypothesized that the increase in WUE associated with lower Gs at elevated [CO2] would reduce the negative impacts of drought on many crops. Despite the large number of CO2-enrichment studies to date, there is relatively little information regarding the effect of elevated [CO2] on stomatal control. Five crop species with active physiological stomatal behavior were grown at ambient (400 ppm) and elevated (2000 ppm) [CO2]. We investigated the relationship between stomatal function, stomatal size, and photosynthetic capacity in the five species, and then assessed the mechanistic effect of elevated [CO2] on photosynthetic physiology, stomatal sensitivity to [CO2] and the effectiveness of stomatal closure to darkness. We observed positive relationships between the speed of stomatal response and the maximum rates of PN and Gs sustained by the plants; indicative of close co-ordination of stomatal behavior and PN. In contrast to previous studies we did not observe a negative relationship between speed of stomatal response and stomatal size. The sensitivity of stomata to [CO2] declined with the ribulose-1,5-bisphosphate limited rate of PN at elevated [CO2]. The effectiveness of stomatal closure was also impaired at high [CO2]. Growth at elevated [CO2] did not affect the performance of photosystem II indicating that high [CO2] had not induced damage to the photosynthetic physiology, and suggesting that photosynthetic control of Gs is either directly impaired at high [CO2], sensing/signaling of environmental change is disrupted or elevated [CO2] causes some physical effect that constrains stomatal opening/closing. This study indicates that while elevated [CO2] may improve the WUE of crops under normal growth conditions, impaired stomatal control may increase the vulnerability of plants to water deficit and high temperatures.

Overwintering evergreen oaks reverse typical relationships between leaf traits in a species spectrum.

The leaf economics spectrum has given us a fundamental understanding of the species variations in leaf variables. Across plant species, tight correlations among leaf mass per area (LMA), mass-based nitrogen (Nm) and photosynthetic rate (Am) and leaf lifespan have been well known as trade-offs in leaf carbon economy. However, the regional or biome-level correlations may not be necessary to correspond with the global-scale analysis. Here, we show that almost all leaf variables in overwintering evergreen oaks in Japan were relatively well included within the evergreen-broadleaved trees in worldwide temperate forests, but Nm was more consistent with that in deciduous broadleaved trees. Contrary to the universal correlations, the correlation between Am and Nm among the evergreen oaks was negative and the correlation between Am and LMA disappeared. The unique performance was due to specific nitrogen allocation within leaves, i.e. the evergreen oaks with later leaf maturation had lower Nm but higher nitrogen allocation to photosynthetic enzymes within leaves, to enhance carbon gain against the delayed leaf maturation and the shortened photosynthetic period due to cold winters. Our data demonstrate that correlations between leaf variables in a local scale are occasionally different from averaged global-scale datasets, because of the constraints in each biome.

Effects of cumulus clouds on microclimate and shoot-level photosynthetic gas exchange in Picea engelmannii and Abies lasiocarpa at treeline, Medicine Bow Mountains, Wyoming, USA

Here we describe the dynamic effects of cumulus clouds on microclimate and shoot-level photosynthetic gas exchange in saplings of two treeline conifer species—Picea engelmannii and Abies lasiocarpa. Measurements were made during both clear-sky and partly cloudy conditions (∼10–70% cumulus cloud cover) throughout the 2012 growing season within an alpine-treeline ecotone. Cumulus clouds generated dynamic fluctuations in photosynthetically active radiation (PAR), higher maximum PAR (>2500μmolm−2s−1), 2–4 fold increases in diffuse PAR, reduced daily mean and cumulative PAR, lower needle temperatures, and reduced leaf-to-air vapor pressure differences relative to clear-sky conditions. Onset of cloud-shade corresponded with declines in photosynthesis, needle temperatures, and evapotranspiration, which were proportional to cloud duration and opacity. Despite increased diffuse light and greater sunlight intensity during cloud-gaps, photosynthesis was never higher on partly cloudy days compared to clear days in either species, during cloud-gaps or cloud-shade. However, reduced transpiration paired with photosynthesis comparable with clear-sky levels during cloud-gaps resulted in greater instantaneous water use efficiency relative to clear-sky measurements. There was no apparent photoinhibition of photosynthesis reflected in gas exchange measurements in response to abrupt and dramatic changes in PAR levels caused by cumulus clouds. We conclude that cumulus clouds reduce instantaneous and daily carbon gain, although lower needle temperatures and associated reductions in transpirational water loss may alleviate daily and seasonal water stress, thereby enhancing carbon gain over the growing season.

Photorespiratory compensation: a driver for biological diversity

This paper reviews how terrestrial plants reduce photorespiration and thus compensate for its inhibitory effects. As shown in the equation φ = (1/Sc/o )O/C, where φ is the ratio of oxygenation to carboxylation, Sc/o is the relative specificity of Rubisco, O is stromal O2 level and C is the stromal CO2 concentration, plants can reduce photorespiration by increasing Sc/o or C, or by reducing O. By far the most effective means of reducing φ is by concentrating CO2, as occurs in C4 and CAM plants, and to a lesser extent in plants using a glycine shuttle to concentrate CO2 into the bundle sheath. Trapping and refixation of photorespired CO2 by a sheath of chloroplasts around the mesophyll cell periphery in C3 plants also enhances C, particularly at low atmospheric CO2. O2 removal is not practical because high energy and protein investment is needed to have more than a negligible effect. Sc/o enhancement provides for modest reductions in φ, but at the potential cost of limiting the kcat of Rubisco. An effective means of decreasing φ and enhancing carbon gain is to lower leaf temperature by reducing absorbance of solar radiation, or where water is abundant, opening stomata. By using a combination of mechanisms, C3 plants can achieve substantial (>30%) reductions in φ. This may have allowed many C3 species to withstand severe competition from C4 plants in low CO2 atmospheres of recent geological time, thereby preserving some of the Earth's floristic diversity that accumulated over millions of years.

Carbon gain, allocation and storage in rhizomes in response to elevated atmospheric carbon dioxide and nutrient supply in a perennial C3 grass, Phalaris arundinacea.

Reed canary grass (Phalaris arundinacea L.) is a fast-growing, perennial, rhizomatous C3 grass considered as a model invasive species for its aggressive behaviour. The same traits make it a candidate for bioenergy feedstock. We tested the following hypotheses: (1) elevated atmospheric [CO2] and nutrient supply enhance photosynthetic carbon acquisition of this fructan-accumulating grass with little or no photosynthetic downregulation; (2) elevated [CO2] promotes carbon allocation to growth when nutrients are sufficient and to fructan storage in rhizomes when nutrients are low. Plants were grown at ambient or elevated (+320μmolmol-1) [CO2], and fertilised using full or one-eighth strength modified Hoagland solution. We investigated leaf photosynthesis, whole-plant water use, biomass allocation, and nitrogen and carbon storage in rhizomes. Elevated [CO2] enhanced light-saturated net CO2 assimilation by 61%. It doubled whole-plant, stem and root biomass in summer. Plants grown in elevated [CO2] had a greater rate of CO2 assimilation at higher [CO2], indicating a shift in photosynthetic apparatus for enhanced carbon gain under elevated [CO2]. The majority of belowground biomass was allocated to rhizomes for storage rather than to roots in both seasons. In autumn, elevated [CO2] increased fructan concentration in rhizomes from 8.1 to 11.7% of biomass when nutrients were low (P=0.023). Our results suggest that elevated [CO2] combined with sufficient nutrients is likely to enhance carbon gain and growth of P. arundinacea, and to increase its productivity and competitiveness in summer. Elevated [CO2] is likely to enhance long-term fructan storage in rhizomes, which may benefit overwintering and vegetative spread.

Net ecosystem CO2 exchange and controlling factors in a steppe—Kobresia meadow on the Tibetan Plateau

Knowledge of seasonal variation of net ecosystem CO2 exchange (NEE) and its biotic and abiotic controllers will further our understanding of carbon cycling process, mechanism and large-scale modelling. Eddy covariance technique was used to measure NEE, biotic and abiotic factors for nearly 3 years in the hinterland alpine steppe—Korbresia meadow grassland on the Tibetan Plateau, the present highest fluxnet station in the world. The main objectives are to investigate dynamics of NEE and its components and to determine the major controlling factors. Maximum carbon assimilation took place in August and maximum carbon loss occurred in November. In June, rainfall amount due to monsoon climate played a great role in grass greening and consequently influenced interannual variation of ecosystem carbon gain. From July through September, monthly NEE presented net carbon assimilation. In other months, ecosystem exhibited carbon loss. In growing season, daytime NEE was mainly controlled by photosynthetically active radiation (PAR). In addition, leaf area index (LAI) interacted with PAR and together modulated NEE rates. Ecosystem respiration was controlled mainly by soil temperature and simultaneously by soil moisture. Q 10 was negatively correlated with soil temperature but positively correlated with soil moisture. Large daily range of air temperature is not necessary to enhance carbon gain. Standard respiration rate at referenced 10°C (R 10) was positively correlated with soil moisture, soil temperature, LAI and aboveground biomass. Rainfall patterns in growing season markedly influenced soil moisture and therefore soil moisture controlled seasonal change of ecosystem respiration. Pulse rainfall in the beginning and at the end of growing season induced great ecosystem respiration and consequently a great amount of carbon was lost. Short growing season and relative low temperature restrained alpine grass vegetation development. The results suggested that LAI be usually in a low level and carbon uptake be relatively low. Rainfall patterns in the growing season and pulse rainfall in the beginning and at end of growing season control ecosystem respiration and consequently influence carbon balance of ecosystem.

Salinity and sea level mediate elevated CO2 effects on C3–C4 plant interactions and tissue nitrogen in a Chesapeake Bay tidal wetland

AbstractElevated atmospheric carbon dioxide concentrations ([CO2]) generally increase plant photosynthesis in C3 species, but not in C4 species, and reduce stomatal conductance in both C3 and C4 plants. In addition, tissue nitrogen concentration ([N]) often fails to keep pace with enhanced carbon gain under elevated CO2, particularly in C3 species. While these responses are well documented in many species, implications for plant growth and nutrient cycling in native ecosystems are not clear. Here we present data on 18 years of measurement of above and belowground biomass, tissue [N] and total standing crop of N for a Scirpus olneyi‐dominated (C3 sedge) community, a Spartina patens‐dominated (C4 grass) community and a C3–C4‐mixed species community exposed to ambient and elevated (ambient +340 ppm) atmospheric [CO2] in natural salinity and sea level conditions of a Chesapeake Bay wetland. Increased biomass production (shoots plus roots) under elevated [CO2] in the S. olneyi‐dominated community was sustained throughout the study, averaging approximately 35%, while no significant effect of elevated [CO2] was found for total biomass in the C4‐dominated community. We found a significant decline in C4 biomass (correlated with rising sea level) and a concomitant increase in C3 biomass in the mixed community. This shift from C4 to C3 was accelerated by the elevated [CO2] treatment. The elevated [CO2] stimulation of total biomass accumulation was greatest during rainy, low salinity years: the average increase above the ambient treatment during the three wettest years (1994, 1996, 2003) was 2.9 t ha−1 but in the three driest years (1995, 1999, 2002), it was 1.2 t ha−1. Elevated [CO2] depressed tissue [N] in both species, but especially in the S. olneyi where the relative depression was positively correlated with salinity and negatively related with the relative enhancement of total biomass production. Thus, the greatest amount of carbon was added to the S. olneyi‐dominated community during years when shoot [N] was reduced the most, suggesting that the availability of N was not the most or even the main limitation to elevated [CO2] stimulation of carbon accumulation in this ecosystem.

Coordinated variation in ecophysiological properties among life stages and tissue types in an invasive perennial forb of semiarid shrub steppe

Semiarid lands of North America are vulnerable to invasion by exotic forbs that frequently have an extended phenology compared with native herbaceous species. We hypothesized that photosynthetic stems, rosette leaves, and cauline leaves of Centaurea maculosa Lam. (knapweed) would differ in ecophysiological specialization and that these differences would enhance plant carbon uptake across seasonal variations in microclimate. Photosynthesis, water relations, and morphological features of C. maculosa were measured under natural variations in temperature and soil water, and under manipulated light environments, in adults compared with seedlings having only rosette leaves. Carbon gain was greater in adults than in seedlings owing to high photosynthesis in cauline and rosette leaves when water was abundant. Otherwise, photosynthesis was relatively lower but persistent through drought in all tissues and ages until senescence. Photosynthesis decreased with water stress in all tissues except stems. Stems comprised up to 36% of photosynthetic area following senescence of rosette leaves during seasonal drought. Seedlings expressed shade acclimation compared with rosette and especially cauline leaves of adults, although adult rosette leaves had flexible photosynthetic light responses. Contrasting ecophysiological specializations of photosynthetic tissues enhance carbon gain of adult C. maculosa as light, water, and temperature vary during its relatively long growth season.

Photosynthetic down‐regulation in Larrea tridentata exposed to elevated atmospheric CO2: interaction with drought under glasshouse and field (FACE) exposure

The photosynthetic response of Larrea tridentata Cav., an evergreen Mojave Desert shrub, to elevated atmospheric CO2 and drought was examined to assist in the understanding of how plants from water‐limited ecosystems will respond to rising CO2. We hypothesized that photosynthetic down‐regulation would disappear during periods of water limitation, and would, therefore, likely be a seasonally transient event. To test this we measured photosynthetic, water relations and fluorescence responses during periods of increased and decreased water availability in two different treatment implementations: (1) from seedlings exposed to 360, 550, and 700 μmol mol–1 CO2 in a glasshouse; and (2) from intact adults exposed to 360 and 550 μmol mol–1 CO2 at the Nevada Desert FACE (Free Air CO2 Enrichment) Facility. FACE and glasshouse well‐watered Larrea significantly down‐regulated photosynthesis at elevated CO2, reducing maximum photosynthetic rate (Amax), carboxylation efficiency (CE), and Rubisco catalytic sites, whereas droughted Larrea showed a differing response depending on treatment technique. Amax and CE were lower in droughted Larrea compared with well‐watered plants, and CO2 had no effect on these reduced photosynthetic parameters. However, Rubisco catalytic sites decreased in droughted Larrea at elevated CO2. Operating Ci increased at elevated CO2 in droughted plants, resulting in greater photosynthetic rates at elevated CO2 as compared with ambient CO2. In well‐watered plants, the changes in operating Ci, CE and Amax resulted in similar photosynthetic rates across CO2 treatments. Our results suggest that drought can diminish photosynthetic down‐regulation to elevated CO2 in Larrea, resulting in seasonally transient patterns of enhanced carbon gain. These results suggest that water status may ultimately control the photosynthetic response of desert systems to rising CO2.

Stomatal dynamics and its importance to carbon gain in two rainforest Piper species : I. VPD effects on the transient stomatal response to lightflecks.

The effects of leaf-air vapor pressure deficit (VPD) on the transient and steady-state stomatal responses to photon flux density (PFD) were evaluated in Piper auritum, a pioneer tree, and Piper aequale, a shade tolerant shrub, that are both native to tropical forests at Los Tuxtlas, Veracruz, México. Under constant high-PFD conditions, the stomata of shade-acclimated plants of both species were sensitive to VPD, exhibiting a nearly uniform decrease in gs as VPD increased. Acclimation of P. auritum to high light increased the stomatal sensitivity to VPD that was sufflcient to cause a reduction in transpiration at high VPD's. At low PFD, where gs was already reduced, there was little additional absolute change with VPD for any species or growth condition. The stomatal response to 8-min duration lightflecks was strongly modulated by VPD and varied between the species and growth light conditions. In P. aequale shade plants, increased VPD had no effect on the extent of stomatal opening but caused the rate of closure after the lightfleck to be faster. Thus, the overall response to a lightfleck changed from hysteretic (faster opening than closure) to symmetric (similar opening and closing rates). Either high or low VPD caused gs not to return to the steady-state value present before the lightfleck. At high VPD the value after was considerably less than the value before whereas at low VPD the opposite occurred. Shade-acclimated plants of P. auritum showed only a small gs response to lightflecks, which was not affected by VPD. Under sunfleck regimes in the understory, the stomatal response of P. aequale at low VPD may function to enhance carbon gain by increasing the induction state. At high VPD, the shift in the response enhances water use efficiency but at the cost of reduced assimilation.

CARBON UPTAKE CHARACTERISTICS IN TWO HIGH ELEVATION POPULATIONS OF THE AQUATIC CAM PLANT ISOETES BOLANDERI (ISOETACAE)

In the aquatic CAM species Isoetes bolanderi, a resident of high elevation lakes in which ambient carbon levels are low and fairly constant throughout the day, CO2 uptake by the leaves generally parallels changes in photosynthetic photon flux density as opposed to changes in ambient CO2 levels. Overnight CO2 uptake by the leaves of I. bolanderi contributed up to 33% of the total daily carbon assimilation from the water column which is typical of aquatic CAM species but low in comparison to the 30–50% seen in the seasonal pool congener Isoetes howellii. Two additional sources of carbon may supplement the 24‐hour period for carbon assimilation conferred by CAM in I. bolanderi: the refixation of respiratory CO2 and carbon acquisition from the interstitial sediment water via roots. CAM appears to play a significant role in enhancing carbon gain in I. bolanderi which dominates the littoral flora of Siesta and Ellery Lakes despite higher carbon uptake rates from the water column found in one associated macrophyte Fontinalis antipyretica.

Carbon Uptake Characteristics in Two High Elevation Populations of the Aquatic Cam Plant Isoetes bolanderi (Isoetacae)

In the aquatic CAM species Isoetes bolanderi, a resident of high elevation lakes in which ambient carbon levels are low and fairly constant throughout the day, CO2 uptake by the leaves generally parallels changes in photosynthetic photon flux density as opposed to changes in ambient CO2 levels. Overnight CO2 uptake by the leaves of I. bolanderi contributed up to 33% of the total daily carbon assimilation from the water column which is typical of aquatic CAM species but low in comparison to the 30–50% seen in the seasonal pool congener Isoetes howellii. Two additional sources of carbon may supplement the 24-hour period for carbon assimilation conferred by CAM in I. bolanderi: the refixation of respiratory CO2 and carbon acquisition from the interstitial sediment water via roots. CAM appears to play a significant role in enhancing carbon gain in I. bolanderi which dominates the littoral flora of Siesta and Ellery Lakes despite higher carbon uptake rates from the water column found in one associated macrophyte Fontinalis antipyretica.

Effect of Leaf Angle and Orientation on Photosynthesis and Water Relations in Silphium terebinthinaceum

Leaf angle and orientation were measured for 217 leaves from two populations of Silphium terebinthinaceum Jacq., a prairie forb with large, unlobed leaves. Seventy-five percent of leaves measured had an angle of deviation from horizontal of more than 60°, and 60% were oriented within 15° of North. Incident Photon Flux Density (PFD), leaf temperature, photosynthesis, stomatal conductance to CO2, internal CO2 concentration, transpiration, and water use efficiency (WUE) were measured on 67 pairs of leaves with the axes oriented in either a North-South (N–S) or East–West (E–W) direction. Leaves with axes oriented N–S intercepted higher levels of PFD during morning and afternoon and exhibited higher diurnal rates of photosynthesis and WUE. Leaf temperature was reduced in N–S leaves during midday as compared to E–W leaves, and was lower in vertical leaves than in those in a horizontal position. Therefore, it was concluded that leaf orientation and verticality enhance carbon gain and minimize water loss—characteristics which may have adaptive significance in a hot, stressful prairie environment.

EFFECT OF LEAF ANGLE AND ORIENTATION ON PHOTOSYNTHESIS AND WATER RELATIONS IN SILPHIUM TEREBINTHINACEUM

Leaf angle and orientation were measured for 217 leaves from two populations of Silphium terebinthinaceum Jacq., a prairie forb with large, unlobed leaves. Seventy‐five percent of leaves measured had an angle of deviation from horizontal of more than 60°, and 60% were oriented within 15° of North. Incident Photon Flux Density (PFD), leaf temperature, photosynthesis, stomatal conductance to CO2, internal CO2 concentration, transpiration, and water use efficiency (WUE) were measured on 67 pairs of leaves with the axes oriented in either a North‐South (N–S) or East–West (E–W) direction. Leaves with axes oriented N–S intercepted higher levels of PFD during morning and afternoon and exhibited higher diurnal rates of photosynthesis and WUE. Leaf temperature was reduced in N–S leaves during midday as compared to E–W leaves, and was lower in vertical leaves than in those in a horizontal position. Therefore, it was concluded that leaf orientation and verticality enhance carbon gain and minimize water loss—characteristics which may have adaptive significance in a hot, stressful prairie environment.

Photosynthetic Utilisation of Lightflecks by Understory Plants

The light environment in forest understories is highly dynamic because the weak shade light is period- ically punctuated by lightflecks lasting from a second or less to tens of minutes. Although present for only a small fraction of the day, these lightflecks can contribute more than two-thirds of the photosynthetically active radiation. Several factots are of importance in determining the capacity of a leaf to utilise lightflecks. Following long low-light periods the induction state of the photosynthetic apparatus is limiting. During induction, 20-60 min may be required before maximum assimilation rates are reached due first to a light activation requirement. of ribulose-1,5-bisphosphate carboxylasel oxygenase and later to the light-induced stomatal opening. Continuous light is not required and induction occurring during a series of lightflecks results in higher carbon gain for later as compared to earlier lightflecks. Post-illumination CO2 fixation resulting from utilisation of metabolite pools built up during the lightfleck can significantly enhance carbon gain during short (5-20 s) lightflecks. The carbon gain of a leaf in response to a lightfleck is a consequence of the limitations imposed by induction state plus the enhancements due to post-illumination CO2 fixation. In the field, this will depend on the frequency and duration of the lightflecks and the duration of the intervening low-light periods.