Photosynthetic Carbon Gain Research Articles

Limitations to photosynthetic carbon gain in timberline Abies lasiocarpa seedlings during prolonged drought

Photosynthesis, water status, and associated physiological parameters were measured in chronically drought-stressed seedlings (5 years of below-average precipitation, 107 cm net deficit) of Abies lasiocarpa (Hook.) Nutt. above (treeline ecotone site, TS) and below (forest site, FS) a Rocky Mountain timberline. In contrast to normal seasonal patterns reported for timberline conifer trees, xylem water potentials were exceptionally low in early summer and remained low for the rest of the summer. Although photosynthesis was not significantly different between the two sites, early season photosynthesis was greater than late-season photosynthesis, especially at FS. Mean daily values of leaf conductance to water vapor (gwv) and transpiration (E) were also low at the beginning of summer (gwv from 0.01 mol·m–2·s–1 to 0.13 mol·m–2·s–1 and E from 0.4 μmol·m–2·s–1 to 2.9 μmol·m–2·s–1) and continued to decrease through summer (an approximate 10-fold decrease in gwv and a 2-fold to 3-fold decrease in E), which resulted in increasing water-use efficiency as summer progressed. Although the slope of instantaneous photosynthesis – intercellular CO2 concentration curves was reduced (lower carboxylation efficiency) from July to September, the relative stomatal limitation to carbon gain was less than 50% over the entire measurement period. Mean daily intercellular CO2 concentrations decreased from near ambient levels (approximately 350–360 ppm) to 290 ppm over the course of summer. Overall, nonstomatal limitations appeared to have the largest impact on photosynthetic carbon gain, although seasonal decreases in leaf conductance and a corresponding depletion of intercellular CO2 indicated that there were also significant stomatal limitations to carbon gain that resulted in a continued regulation of greater water use efficiency.

Cytokinin Import Rate as a Signal for Photosynthetic Acclimation to Canopy Light Gradients

Plants growing in dense canopies are exposed to vertical light gradients and show photosynthetic acclimation at the whole-plant level, resulting in efficient photosynthetic carbon gain. We studied the role of cytokinins transported through the transpiration stream as one of probably multiple signals for photosynthetic acclimation to light gradients using both tobacco (Nicotiana tabacum) and Arabidopsis (Arabidopsis thaliana). We show that substantial variation in leaf transpiration parallels the light gradient in tobacco canopies and experimental reduction of the transpiration rate of a leaf, independent of light, is sufficient to reduce photosynthetic capacity in both species, as well as transcript levels of the small subunit of Rubisco (rbcS) gene in Arabidopsis. Mass spectrometric analysis of xylem sap collected from intact, transpiring tobacco plants revealed that shaded leaves import less cytokinin than leaves exposed to high light. In Arabidopsis, reduced transpiration rate of a leaf in the light is associated with lower cytokinin concentrations, including the bioactive trans-zeatin and trans-zeatin riboside, as well as reduced expression of the cytokinin-responsive genes ARR7 and ARR16. External application of cytokinin to shaded leaves rescued multiple shade effects, including rbcS transcript levels in both species, as did locally induced cytokinin overproduction in transgenic tobacco plants. From these data, we conclude that light gradients over the foliage of a plant result in reduced cytokinin activity in shaded leaves as a consequence of reduced import through the xylem and that cytokinin is involved in the regulation of whole-plant photosynthetic acclimation to light gradients in canopies.

Increased air temperature during simulated autumn conditions does not increase photosynthetic carbon gain but affects the dissipation of excess energy in seedlings of the evergreen conifer Jack pine.

Temperature and daylength act as environmental signals that determine the length of the growing season in boreal evergreen conifers. Climate change might affect the seasonal development of these trees, as they will experience naturally decreasing daylength during autumn, while at the same time warmer air temperature will maintain photosynthesis and respiration. We characterized the down-regulation of photosynthetic gas exchange and the mechanisms involved in the dissipation of energy in Jack pine (Pinus banksiana) in controlled environments during a simulated summer-autumn transition under natural conditions and conditions with altered air temperature and photoperiod. Using a factorial design, we dissected the effects of daylength and temperature. Control plants were grown at either warm summer conditions with 16-h photoperiod and 22 degrees C or conditions representing a cool autumn with 8 h/7 degrees C. To assess the impact of photoperiod and temperature on photosynthesis and energy dissipation, plants were also grown under either cold summer (16-h photoperiod/7 degrees C) or warm autumn conditions (8-h photoperiod/22 degrees C). Photosynthetic gas exchange was affected by both daylength and temperature. Assimilation and respiration rates under warm autumn conditions were only about one-half of the summer values but were similar to values obtained for cold summer and natural autumn treatments. In contrast, photosynthetic efficiency was largely determined by temperature but not by daylength. Plants of different treatments followed different strategies for dissipating excess energy. Whereas in the warm summer treatment safe dissipation of excess energy was facilitated via zeaxanthin, in all other treatments dissipation of excess energy was facilitated predominantly via increased aggregation of the light-harvesting complex of photosystem II. These differences were accompanied by a lower deepoxidation state and larger amounts of beta-carotene in the warm autumn treatment as well as by changes in the abundance of thylakoid membrane proteins compared to the summer condition. We conclude that photoperiod control of dormancy in Jack pine appears to negate any potential for an increased carbon gain associated with higher temperatures during the autumn season.

Nighttime Stomatal Conductance and Transpiration in C3 and C4 Plants

Incomplete stomatal closure during the night is observed in a diverse range of C3 and C4 species ([Fig. 1][1] ; Supplemental Table S1) and can lead to substantial nighttime transpirational water loss. Although water loss is an inevitable consequence of stomatal opening for photosynthetic carbon gain

Significant transpirational water loss occurs throughout the night in field-grown tomato.

Incomplete stomatal closure at night can result in substantial water loss at times when photosynthetic carbon gain is not occurring in C3 and C4 plant species. To investigate the magnitude of nighttime water loss for a crop species in the field, measurements of nighttime water loss by tomato (Lycopersicon esculentum Mill. cv. Heinz 8892) were made by three methods: a field-scale lysimeter and two leaf-level instruments, an automated viscous flow porometer and a portable photosynthesis system. The portable photosynthesis system indicated nighttime transpiration of 10% of maximal daytime transpiration and the viscous flow porometer demonstrated partially open stomata. Integrated crop water loss during the dark, non-photosynthetic hours measured on the lysimeter was 3-10.8% of total daily water loss. In the glasshouse, a survey of closely related wild and cultivated tomato species showed that under ambient conditions nighttime transpiration varied within and among species and was 8-33% of maximal daytime transpiration. Implications of such a substantial fraction of total daily crop water use occurring during the night are significant in agronomic, environmental, and economic terms. Further, variation within and among species in nighttime water loss has implications for breeding to improve crop water use efficiency.

Temporal and light-based changes in carbon uptake and storage in the spring ephemeral Podophyllum peltatum (Berberidaceae)

The single leaf of the spring ephemeral Podophyllum peltatum (Berberidaceae), emerges in the spring, when forest over-story leaf area index (LAI) is low (∼0.8) and senesces approximately 12 weeks later when LAI is high (∼4.0). This study characterized (i) seasonal changes in gas exchange characteristics (maximum light-saturated photosynthetic rate, foliar dark respiration rate, apparent quantum efficiency, light saturation point and light compensation point) and carbohydrate storage in field populations and (ii) examined the hypothesis that observed seasonal changes in photosynthetic characteristics represent photosynthetic acclimation to maximize photosynthetic carbon gain as light availability declined. In field populations, as LAI increased, total chlorophyll concentration increased while declines occurred in all gas exchange characteristics except apparent quantum efficiency. Over the same period, soluble sugar content of the rhizome declined and starch content increased rapidly during the first 4 weeks after which it stabilized. Foliar tissue exhibited little change in carbohydrate pool over the season. Laboratory populations grown at high and low light levels differed in all gas exchange parameters except light saturation point, but illustrated similar seasonal changes in gas exchange characteristics to both each other and the field population. The similarity of temporal changes in gas exchange characteristics of field (seasonal decline in light availability) and laboratory (no seasonal decline in light availability) suggests that P. peltatum possesses limited ability to photosynthetically acclimate to declining light availability. It appears that seasonal carbon gain in P. peltatum may be primarily controlled by leaf lifespan and the fraction of lifespan occurring under high illumination conditions early in the season.

Linking physiological processes with mangrove forest structure: phosphorus deficiency limits canopy development, hydraulic conductivity and photosynthetic carbon gain in dwarf Rhizophora mangle

Spatial gradients in mangrove tree height in barrier islands of Belize are associated with nutrient deficiency and sustained flooding in the absence of a salinity gradient. While nutrient deficiency is likely to affect many parameters, here we show that addition of phosphorus (P) to dwarf mangroves stimulated increases in diameters of xylem vessels, area of conductive xylem tissue and leaf area index (LAI) of the canopy. These changes in structure were consistent with related changes in function, as addition of P also increased hydraulic conductivity (Ks), stomatal conductance and photosynthetic assimilation rates to the same levels measured in taller trees fringing the seaward margin of the mangrove. Increased xylem vessel size and corresponding enhancements in stem hydraulic conductivity in P fertilized dwarf trees came at the cost of enhanced mid-day loss of hydraulic conductivity and was associated with decreased assimilation rates in the afternoon. Analysis of trait plasticity identifies hydraulic properties of trees as more plastic than those of leaf structural and physiological characteristics, implying that hydraulic properties are key in controlling growth in mangroves. Alleviation of P deficiency, which released trees from hydraulic limitations, reduced the structural and functional distinctions between dwarf and taller fringing tree forms of Rhizophora mangle.

Photosynthesis during an Episodic Drought in Abies lasiocarpa and Picea engelmannii across an Alpine Treeline

Effects of abiotic factors on daily and annual photosynthetic carbon gain were evaluated in Abies lasiocarpa and Picea engelmannii at three sites across an alpine treeline ecotone in the Medicine Bow Mountains of southeastern Wyoming (U.S.A.). In addition, the year 2001–2002 was characterized as an episodic drought (seventh driest year since 1895), including a winter that generated only 45% of normal snowpack. Both species had approximately 50% lower xylem water potentials and photosynthesis compared to previous studies for the same species and locale. In A. lasiocarpa, estimated total photosynthesis for the measurement period (A tot) was greatest (28.7 mol m−2) at the mid-ecotone site (∼3198 m), followed by the alpine site (24.6 mol m−2) at ∼3286 m, and then the forest site (19.4 mol m−2) below timberline (∼2965 m). Similar results occurred in P. engelmannii (17.6, 23.4, and 25.3 mol m−2, respectively). These differences appeared to be most influenced by stomatal rather than non-stomatal effects based on comparisons of photosynthesis, leaf conductance, and internal CO2 concentrations through summer. Although photosynthesis over the summer appeared limited primarily by annual water limitation, the two higher elevation sites had significantly greater values that were associated with microclimatic differences in sunlight incidence and, possibly, temperature.

Refugial forests of the southern Appalachians: photosynthesis and survival in current-year Abies fraseri seedlings

Fraser fir (Abies fraseri (Pursh) Poiret) is an endemic, high-elevation conifer confined to six relict mountaintop communities in the southern Appalachian Mountains, USA. High adult mortality has occurred over the past 50 years, possibly the result of an introduced insect (Adelges piceae Ratzeburg), air pollution, or both. Knowledge of the mechanisms of and limitations to seedling establishment may allow reestablisment and perpetuation of this unique community type, notwithstanding global climate change. We monitored seedling emergence and mortality in relation to photosynthetic performance and water relations in microsites differing in canopy openness (sunlight exposure) over the summer of 2004. Abundance of cotyledonous seedlings in early summer was 2.3 times greater (849 versus 366 seedlings m(-2)) in microsites with lower sky exposure (greater canopy closure) than in microsites with greater sky exposure (greater canopy openness). In contrast, late-season abundance and survival were greater in areas beneath more open canopies than in areas beneath less open canopies (3.3 times and 11.7 times greater, respectively). However, newly emerged seedling survival in a completely open site (no overhead canopy) was zero, despite an initial density of 124 seedlings m(-2). Seedling water status was similar in open- and closed-canopy sites (-0.52 and -0.74 MPa, respectively). Photosynthetic carbon gain was higher in newly emerged seedlings at open canopy than at closed canopy sites, especially during early morning. Based on photosynthetic light response curves and measured sunlight regimes, seedlings in open canopy sites were estimated to assimilate 3.3-4.5 times more carbon than seedlings at closed sites. Reductions in carbon gain of closed-site seedlings, primarily a result of limited sunlight, corresponded to substantial increases in seedling mortality (98 versus 79% in open canopy sites). Thus, sunlight exposure, which reflects overstory canopy structure, appears to be an important factor influencing newly emerged seedling survival and distribution.

Photosynthetic and structural characteristics of canopy and shrub trees in a cool-temperate deciduous broadleaved forest: Implication to the ecosystem carbon gain

To reveal the seasonal change of leaf ecophysiological and canopy characteristics and to evaluate the functional role of canopy and shrub tree species in forest CO 2 uptake, we measured forest canopy leaf area index (LAI) using a hemispherical canopy photography technique, leaf CO 2 gas exchange and shoot architecture for canopy ( Betula ermanii and Quercus crispula) and shrub ( Hydrangea paniculata and Viburnum furcatum) tree species in a deciduous broadleaved forest in a cool-temperate region in central Japan. Canopy LAI and photosynthetic capacity of canopy tree leaves increased rapidly with leaf expansion. LAI reached its maximum in early summer but photosynthetic capacity reached its maximum in late summer. Development of photosynthetic capacity was dependent on the changes of leaf mass per area and leaf chlorophyll content (evaluated by SPAD). The seasonal maximum photosynthetic capacity of the leaves at the forest canopy top ( B. ermanii and sun leaves of Q. crispula) was about more than double of the leaves in the shrub layer ( H. paniculata, shade leaves of Q. crispula and V. furcatum). Light interception and photosynthetic carbon gain at a shoot level were simulated under three air temperature conditions by a three-dimensional canopy photosynthesis model (Y-plant) involving the combined leaf photosynthesis and stomatal conductance responses and shoot architecture. Results showed that (1) calculations without considering the heterogeneous light distribution in a foliage made by geometrical feature of plants would overestimate the photosynthetic carbon gain by +40% even at the canopy surface, and (2) the steep leaf angle in B. ermanii avoided midday depression of photosynthesis while the rather horizontal leaves in Q. crispula received excess light and heat load which led larger midday depression of photosynthesis. In addition to the large capacity of photosynthetic productivity of the canopy top foliage, our model also suggests the functional role of shrub species in forest ecosystem carbon gain, due to their high photosynthetic utilization efficiency of low light incidence available in the forest understory.

Non-destructive estimation of lateral root distribution in an aridland perennial

Estimation of root distributions in natural systems remains challenging due to the difficulties in excavation and easy breakage of fine roots. Identifying lateral fine root distribution is necessary to determine the potential exploitation of spatially and temporally variable nutrient supplies that characterize most arid ecosystems. We estimated this potential by taking field measurements of lateral root distribution of the small herbaceous perennial Cryptantha flava (A. Nels.) Payson using 15N-enriched nutrient solutions wicked into the soil at various distances from study plants. Leaves were subsequently harvested from these plants and analyzed for N isotopic ratios. C. flava plants were capable of N uptake at distances of greater than 1.0 m from the outer edge of their aboveground canopy. The considerable lateral root neighborhood area of C. flava increases the amount of spatially variable N that is exploitable in these low-N soils. The ability to acquire spatially variable N and rapidly translate N uptake into photosynthetic carbon gain are traits that aid C. flava in maintaining its position as a successful subordinate competitor in a community dominated by larger, woody perennials.

Diversity of algal endosymbionts (zooxanthellae) in octocorals: the roles of geography and host relationships

The presence, genetic identity and diversity of algal endosymbionts (Symbiodinium) in 114 species from 69 genera (20 families) of octocorals from the Great Barrier Reef (GBR), the far eastern Pacific (EP) and the Caribbean was examined, and patterns of the octocoral-algal symbiosis were compared with patterns in the host phylogeny. Genetic analyses of the zooxanthellae were based on ribosomal DNA internal transcribed spacer 1 (ITS1) region. In the GBR samples, Symbiodinium clades A and G were encountered with A and G being rare. Clade B zooxanthellae have been previously reported from a GBR octocoral, but are also rare in octocorals from this region. Symbiodinium G has so far only been found in Foraminifera, but is rare in these organisms. In the Caribbean samples, only Symbiodinium clades B and C are present. Hence, Symbiodinium diversity at the level of phylogenetic clades is lower in octocorals from the Caribbean compared to those from the GBR. However, an unprecedented level of ITS1 diversity was observed within individual colonies of some Caribbean gorgonians, implying either that these simultaneously harbour multiple strains of clade B zooxanthellae, or that ITS1 heterogeneity exists within the genomes of some zooxanthellae. Intracladal diversity based on ITS should therefore be interpreted with caution, especially in cases where no independent evidence exists to support distinctiveness, such as ecological distribution or physiological characteristics. All samples from EP are azooxanthellate. Three unrelated GBR taxa that are described in the literature as azooxanthellate (Junceella fragilis, Euplexaura nuttingi and Stereonephthya sp. 1) contain clade G zooxanthellae, and their symbiotic association with zooxanthellae was confirmed by histology. These corals are pale in colour, whereas related azooxanthellate species are brightly coloured. The evolutionary loss or gain of zooxanthellae may have altered the light sensitivity of the host tissues, requiring the animals to adopt or reduce pigmentation. Finally, we superimposed patterns of the octocoral-algal symbiosis onto a molecular phylogeny of the host. The data show that many losses/gains of endosymbiosis have occurred during the evolution of octocorals. The ancestral state (azooxanthellate or zooxanthellate) in octocorals remains unclear, but the data suggest that on an evolutionary timescale octocorals can switch more easily between mixotrophy and heterotrophy compared to scleractinian corals, which coincides with a low reliance on photosynthetic carbon gain in the former group of organisms.

Enhanced photosynthesis and flower production in a sagebrush morphotype associated with animal burrows

Two morphotypes of the evergreen shrub Artemisia tridentata Nutt. ssp. wyomingensis occur in the Shirley Basin of central Wyoming (USA), one of which was associated exclusively with Mima-like mounds generated by animal burrowing activity. Measured on a particularly dry year according to a 34-year precipitation record, plants growing on mounds (M) versus inter-mound locations (IM)were taller with greater leaf biomass and leaf area per unit ground area, and had over 90% of all inflorescences. As a result, the landscape consists of a patchy distribution of reproductive islands (~ 20-40 m-2 in size) separated by a mean distance of ~ 30 m. In addition, greater photosynthesis per unit leaf area occurred for M plants when ephemeral leaves dominated total leaf area in spring and early summer, as well as during short time periods (< 3 days) following sporadic rainfall events in summer when only perennial leaves were present. As a result, estimated total annual carbon gain was 41% greater for M plants from May to mid-June, but was not significantly different from IM plants for the remainder of the season, resulting in a total summer carbon gain that was 14% greater in M plants. Stomatal and nonstomatal conductances to CO2 uptake were also greater for the ephemeral leaves of M plants, along with lower internal CO2 concentrations (193 ± 4 μl l-1 vs. 209 ± 8 μl-1, respectively). M plants also maintained higher xylem water potentials throughout most of the growth season (−1.1 ± 0.1 SD MPa in May, declining to −4.4 ± 0.3 SD MPa in August), along with higher water use efficiencies (photosynthesis/transpiration). M and IM soils did not differ significantly in total organic or nitrate contents, although leaf nitrogen content was higher in M plants when photosynthesis was also greater. Photosynthesis in M plants also responded more positively to afternoon showers greater than about 7 mm compared to IM plants. Thus, improved water and nutrient relations was associated with enhanced photosynthetic carbon gain in M plants, enabling greater flower production. Moreover, morphotypic plasticity coupled with the effects of animal burrows may have substantially increased sexual reproductive success in A. t. wyomingensis.

Physiological and ecological significance of sunflecks for dipterocarp seedlings

Irradiance is highly dynamic in many plant canopies. Photosynthesis during sunflecks provides 10-90% of daily carbon gain. The survivorship of tree seedlings in the deeply shaded understorey of tropical rain forests is limited by their ability to maintain a positive carbon balance. Dipterocarp seedlings from the SE Asian rain forest were used as a model system to test novel aspects of the physiological and ecological significance of sunflecks. First, understorey seedlings experienced leaf temperatures up to 38 degrees C in association with sunflecks. Under controlled environment conditions, the inhibition of carbon gain at 38 degrees C, compared with 28 degrees C, was significantly greater during a sequence of sunflecks (-59%), than under uniform irradiance (-40%), providing the same total photosynthetic photon flux density (PPFD). Second, the relative enhancement effects of elevated [CO2] were greater under sunflecks (growth +60%, carbon gain +89%), compared with uniform irradiance (growth +25%, carbon gain +59%), supplying the same daily PPFD. Third, seedling growth rates in the forest understorey were 4-fold greater under a dynamic irradiance treatment characterized by long flecks, compared with a regime of short flecks. Therefore, stresses associated with dynamic irradiance may constrain photosynthetic carbon gain. Additionally, seedling photosynthesis and growth may be more responsive to interactions with abiotic factors, including future changes in climate, than previously estimated. The sensitivity of seedling growth to varying patterns of dynamic irradiance, and the increased likelihood of species-specific responses through interactions with environmental factors, indicates the potential for sunflecks to influence regeneration processes, and hence forest structure and composition.

Photosynthetic rates and partitioning of absorbed light energy in photoinhibited leaves

Parameters for the evaluation of the effects of photoinhibition on photosynthetic carbon gain were studied in Chenopodium album leaves. The light‐response curve of photosynthetic rate was determined at 36 Pa CO2 partial pressure and fitted by a non‐rectangular hyperbola. Both the initial slope of the curve and the light‐saturated rate decreased in photoinhibited leaves, although the decrease in the latter was small. The convexity of the curve was also smaller in photoinhibited leaves. The capacities of ribulose‐1,5‐bisphosphate carboxylation (Vcmax) and electron transport (Jmax) were estimated from the CO2‐response curves. Vcmax and Jmax decreased similarly with increasing photoinhibition. Energy partitioning in photosystem II (PSII) was estimated using chlorophyll fluorescence parameters. The fraction of energy that was consumed by photochemistry decreased with increasing photoinhibition. However, an increase in inactive PSII, decreasing energy partitioning to active PSII, relaxed the excitation pressure in PSII, and led to a reduction in the fraction of excess energy that was neither consumed by photochemistry nor dissipated as heat.

Leaf Orientation, Incident Sunlight, and Photosynthesis in the Alpine Species Suassurea superba and Gentiana straminea on the Qinghai-Tibet Plateau

The extremely high level of solar radiation on the Qinghai-Tibet Plateau may induce photoinhibition and thus limit leaf carbon gain. To assess the effect of high light, we examined gas exchange and chlorophyll fluorescence for two species differing in light interception: the prostrate Saussurea superba and the erect-leaved Gentiana straminea. In controlled conditions with favorable water and temperature, neither species showed apparent photoinhibition in gas exchange measurements. In natural environment, however, their photosynthetic rate decreased remarkably at high light. Photosynthesis depression was aggravated under high leaf temperature or soil water stress. Relative stomatal limitation was much higher in S. superba than in G. straminea and it remarkably increased in the later species at midday when soil was dry. F v /F m as an indicator for photoinhibition was generally higher in S. superba than in the other species. F v /F m decreased significantly under high light at midday in both species, even when soil moisture was high. F 0 linearly elevated with the increment of leaf temperature in G. straminea, but remained almost constant in S. superba. Electron transport rate (ETR) increased with photosynthetically active photon flux density (PPFD) in S. superba, but declined when PPFD was high than about 1000 μmol m−2 s−1 in G. straminea. Compared to favorable environment, the estimated daily leaf carbon gain at PPFD above 800 μmol m−2 s−1 was reduced by 32% in S. superba and by 17% in G. straminea when soil was moist, and by 43% and 53%, respectively, when soil was dry. Our results suggest that the high radiation induces photoinhibition and significantly limits photosynthetic carbon gain, and the limitation may further increase at higher temperature and in dry soil.

Direct and indirect climate change effects on photosynthesis and transpiration.

Climate change affects plants in many different ways. Increasing CO(2) concentration can increase photosynthetic rates. This is especially pronounced for C(3) plants, at high temperatures and under water-limited conditions. Increasing temperature also affects photosynthesis, but plants have a considerable ability to adapt to their growth conditions and can function even at extremely high temperatures, provided adequate water is available. Temperature optima differ between species and growth conditions, and are higher in elevated atmospheric CO(2). With increasing temperature, vapour pressure deficits of the air may increase, with a concomitant increase in the transpiration rate from plant canopies. However, if stomata close in response to increasing CO(2) concentration, or if there is a reduction in the diurnal temperature range, then transpiration rates may even decrease. Soil organic matter decomposition rates are likely to be stimulated by higher temperatures, so that nutrients can be more readily mineralised and made available to plants. This is likely to increase photosynthetic carbon gain in nutrient-limited systems. All the factors listed above interact strongly so that, for different combinations of increases in temperature and CO(2) concentration, and for systems in different climatic regions and primarily affected by water or nutrient limitations, photosynthesis must be expected to respond differently to the same climatic changes.

Synchronization of metabolic processes in plants with Crassulacean acid metabolism.

In plants with Crassulacean acid metabolism, a diel separation of carboxylation processes mediated by phosphoenolpyruvate carboxylase (PEPC) and Rubisco optimizes photosynthetic performance and carbon gain in potentially limiting environments. This review considers the mechanisms that synchronize the supply and demand for carbon whilst maintaining photosynthetic plasticity over the 24 h CAM cycle. The circadian clock plays a central role in controlling many of the metabolic, transport and physiological components of CAM. The level of control exerted by the clock can range from transcriptional through to post-translational regulation, depending on the genes, proteins, and even plant species under consideration. A further layer of control is provided by metabolites, including organic acids and carbohydrates, which show substantial reciprocal fluctuations in content over the diel cycle. Mechanisms responsible for the sensing of metabolite contents are discussed, together with signalling requirements for the co-ordination of carbon fluxes. Evolutionary implications are considered in terms of how circadian and metabolic control of the CAM cycle may have been derived from C3 plants.

Low temperature effects on photosynthesis and growth of grapevine

ABSTRACTGrowth and photosynthesis of grapevine (Vitis vinifera L.) planted on two sloping cool climate vineyards were measured during the early growth season. At both vineyards, a small difference in mean minimum air temperature (1–3 °C) between two microsites accumulated over time, producing differences in shoot growth rate. The growth rates of the warmer (upper) microsite were 34–63% higher than the cooler (lower) site. Photosynthesis measurements of both east and west canopy sides revealed that the difference in carbon gain between the warmer and cooler microsites was due to low temperatures restricting the photosynthetic contribution of east‐facing leaves. East‐facing leaves at the warmer microsite experienced less time at suboptimal temperature while being exposed to high irradiance, contributing to an average 10% greater net carbon gain compared to the east‐facing leaves at the cooler microsite. This chilling‐induced reduction in photosynthesis was not due to net photo‐inhibition. Further analysis revealed that CO2‐ and light‐saturated photosynthesis of grapevines was restricted by stomatal closure from 15 to 25 °C and by a limitation of RuBP regeneration and/or end‐product limitation from 5 to 15 °C. Changes in photosynthetic carboxylation efficiency implied that Rubisco activity may also play a regulatory role at all temperatures. This restriction of total photosynthetic carbon gain is proposed to be a major contributor to the temperature dependence of growth rate at both vineyards during the early season growth period.

Abiotic factors limiting photosynthesis in Abies lasiocarpa and Picea engelmannii seedlings below and above the alpine timberline.

Most research on the occurrence and stability of alpine timberlines has focused on correlations between adult tree growth and mean temperatures rather than on specific mechanisms. Timberline migration to higher altitude is dependent on new seedling establishment in the tree-line ecotone; however, reductions in photosynthetic carbon gain in establishing seedlings have previously been interpreted solely in terms of decreased seedling survival. Our objective was to evaluate the impact of abiotic factors (temperature, light and water) on photosynthetic carbon gain in young seedlings of the two dominant conifer tree species occurring naturally above (tree-line ecotone site, TS) and below (forest site, FS) a Rocky Mountain timberline in southeastern Wyoming, USA. Coincidentally, measurements were made during an unusually dry summer. Mean daily photosynthesis in seedlings of both Abies lasiocarpa (Hook.) Nutt. (subalpine fir) and Picea engelmannii Parry ex Engelm. (Engelmann spruce) was less at TS than at FS (19 and 29%, respectively). Minimum nighttime temperatures below 2 degrees C were more frequent at TS than at FS and were associated with reduced maximum photosynthesis the following day. Low midday water potentials were associated with a reduction in carbon gain at both sites early in the season, prior to snowmelt, as well as late in the season when soils began to dry. However, the lower photosynthetic rates at TS than at FS appeared to be unrelated to seedling water status because seedlings at both sites had similar xylem pressure potentials. Solar irradiance was highly variable at both sites as a result of uneven shading by neighboring trees, although this variation was substantially reduced on cloudy days (44% of all days observed). Compared with sunny days, cloudy days resulted in greater integrated daily carbon gain at both sites (41% increase at TS and 69% increase at FS), based on a simulated photosynthesis model. Photosynthetic responses to temperature, sunlight and water suggest that variable solar irradiance and nighttime temperatures were major abiotic factors limiting photosynthetic carbon acquisition in these young seedlings, especially for seedlings growing in the tree-line ecotone.