Water relations and carbon gain are closely related to cushion size in the moss Grimmia pulvinata

The present study of structural and physiological changes during the development of the cushion moss, Grimmia pulvinata , quantifies the size-dependence of various parameters of water relations such as changes in surface: volume ratio ( S/V ) or water loss rates, and also measures net CO₂ gas exchange in the light and the dark. Larger cushions had lower S/V values than smaller ones and featured lower rates of area-based evapotranspiration, owing to higher boundary-layer resistance, but did not differ in relative water storage capacity (expressed as a percentage of d. wt). In combination, this leads to considerably longer hydration periods in larger cushions. By contrast, CO₂ gas-exchange parameters were negatively correlated with size : larger cushions showed significantly lower (mass-based) rates of net photosynthesis and dark respiration. Using these data, we estimated carbon budgets during a drying cycle as a function of cushion size. When including alternations of dark and light periods, the relationship proved to be rather complicated. Depending on the time of hydration, net carbon budgets not only varied quantitatively with size but sometimes took on both positive and negative values depending on cushion size. We conclude that neglecting plant size can lead to unrepeatable or even misleading results in comparative ecophysiological studies, and therefore urge for adequate attention to be paid to size in these studies.     

Carbon gains by desiccation-tolerant plants at elevated CO2

1. There have been no reports of the long-term responses of the desiccation-tolerant (DT) plants to elevated CO₂. Xerophyta scabrida is a DT woody shrub, which loses chlorophylls and thylakoids during desiccation: a so-called poikilochlorophyllous desiccation-tolerant species (PDT). When the leaves of X. scabria are allowed to desiccate, the species shows many of the normal features of (P)DT plants.
2. However, the duration of photosynthesis in X. scabria is prolonged by 300% when the measurements are made at 700 as opposed to 350p.p.m. CO₂. The implication is that the carboxylating enzymes must still have been active at this time to enable appreciable photosynthetic activity. This response could have far-reaching implications for the success of such species in a future climate.
3. Lichens and mosses, representing the homoiochlorophyllous DTs (HDT), retain their chlorophyll content and photosynthetic apparatus during desiccation. We show the desiccation responses of two common HDT species ( Cladonia convoluta and Tortula ruralis ) to elevated CO₂ for comparison. Both HDT species showed increased net CO₂ uptake in the material grown at high CO₂ by more than 30% in moss and by more than 50% in lichen. It is concluded that desiccation-tolerant plants will be among the main beneficiaries of a high CO₂ future.     

Leaf stomatal responses to vapour pressure deficit under current and CO2-enriched atmosphere explained by the economics of gas exchange

Using the economics of gas exchange, early studies derived an expression of stomatal conductance ( g ) assuming that water cost per unit carbon is constant as the daily loss of water in transpiration ( f _e) is minimized for a given gain in photosynthesis ( f _c). Other studies reached identical results, yet assumed different forms for the underlying functions and defined the daily cost parameter as carbon cost per unit water. We demonstrated that the solution can be recovered when optimization is formulated at time scales commensurate with the response time of g to environmental stimuli. The optimization theory produced three emergent gas exchange responses that are consistent with observed behaviour: (1) the sensitivity of g to vapour pressure deficit ( D ) is similar to that obtained from a previous synthesis of more than 40 species showing g to scale as 1 −  m  log( D ), where m   ∈  [0.5,0.6], (2) the theory is consistent with the onset of an apparent 'feed-forward' mechanism in g , and (3) the emergent non-linear relationship between the ratio of intercellular to atmospheric [CO₂] ( c _i/ c _a) and D agrees with the results available on this response. We extended the theory to diagnosing experimental results on the sensitivity of g to D under varying c _a.     

Effects of boundary layer conductance on substomatal pressures of carbon dioxide

Abstract. Gas exchange measurements were made on single leaves of three C₃ and one C₄ species at air speeds of 0.4 and 4.0 m s⁻¹ to determine if boundary layer conductance substantially affected the substomatal pressure of carbon dioxide. Boundary layer conductances to water vapour were 0.4 to 0.5 mol m⁻² s⁻¹ at the lower air speed, and 1.2 to 1.5 mol m⁻² s⁻¹ at the higher air speed. Substomatal carbon dioxide pressures were about 5 Pa lower at low boundary layer conductance in the C₃ species, and about 3 Pa lower in the C₄ species when measurements were made at high and moderate photosynthetic photon flux densities. No evidence of stomatal adjustment to altered boundary layer conductance was found. Photosynthetic rates at high photon flux densities were reduced by about 20% at the low air speed in the C₃ species. The commonly reported values of substomatal carbon dioxide pressure for C₃ and C₄ species were found to occur only when measurements were made at the higher air speed.     

Stomatal and photosynthetic responses to shade in sorghum, soybean and eastern gamagrass

We studied photosynthetic and stomatal responses of grain sorghum ( Sorghum bicolor [L.] Moench cv. Pioneer 8500), soybean ( Glycine max L. cv. Flyer) and eastern gamagrass ( Tripsacum dactyloides L.) during experimental sun and shade periods simulating summer cloud cover. Leaf gas exchange measurements of field plants showed that short-term (5 min) shading of leaves to 300–400 μmol m⁻² s⁻¹ photosynthetic photon flux density reduced photosynthesis, leaf temperature, stomatal conductance, transpiration and water use efficiency and increased intercellular CO₂ partial pressure. In all species, photosynthetic recovery was delayed when leaves were reilluminated, apparently by stomatal closure. The strongest stomatal response was in soybean. Photosynthetic recovery was studied further with soybeans grown indoors (maximum photosynthetic photon flux density 1 200 μmol m⁻² s⁻¹). Plants grown indoors had responses to shade similar to those of field plants, except for brief nonstomatal limitation immediately after reillumination. These responses indicated the importance of the light environment during leaf development on assimilation responses to variable light, and suggested different limitations on carbon assimilation in different parts of the soybean canopy. Photosynthetic oxygen evolution recovered immediately upon reillumination, indicating that the light reactions did not limit soybean photosynthetic recovery. While shade periods caused stomatal closure and reduced carbon gain and water loss in all species, the consequences for carbon gain/water loss were greatest in soybean. The occurrence of stomatal closure in all three species may arise from their shared phenologies and herbaceous growth forms.     

Leaf Phenology and Seasonal Carbon Gain in the Invasive Plant, Bunias orientalis L.

Abstract: In two potentially competing herbaceous plants, the invasive Bunias orientalis L. (Brassicaceae) and the native Picris hieracioides L. (Asteraceae), seasonal changes in leaf CO₂ gas exchange and plant growth were studied over an entire growing season from February 1998 to December 1998 in two experimental fields. The study was motivated by the hypothesis that pre-adaptive phenological displacement of alien species relative to the native flora may be an important reason for the observed expansion of B. orientalis in central Europe. We quantified the importance of phenological differences for annual carbon gain in both species by estimating total leaf carbon gain from the results of leaf CO₂ exchange and changes in plant leaf area. Bunias orientalis achieved almost half of its annual carbon gain in the time between early September and December, when competition for light by other species, like P. hieracioides , is low. Our quantitative approach corroborates the notion that the phenological shift of a relatively poor competitor, such as B. orientalis , could be of great importance for the success as an invasive species.     

Physiological and isotopic (δ13C and δ18O) responses of three tropical tree species to water and nutrient availability

Water-use efficiency and stable isotope composition were studied in three tropical tree species. Seedlings of Tectona grandis , Swietenia macrophylla and Platymiscium pinnatum were grown at either high or low water supply, and with or without added fertilizer. These three species previously exhibited low, intermediate and high whole-plant water-use efficiency ( TE ) when grown at high water supply in unfertilized soil. Responses of TE to water and nutrient availability varied among species. The TE was calculated as experiment-long dry matter production divided by cumulative water use. Species-specific offsets were observed in relationships between TE and whole-plant ¹³C discrimination (Δ¹³C_p). These offsets could be attributed to a breakdown in the relationship between Δ¹³C_p and the ratio of intercellular to ambient CO₂ partial pressures ( c _i/ c _a) in P. pinnatum , and to variation among species in the leaf-to-air vapour pressure difference ( v ). Thus, a plot of v · TE against c _i/ c _a showed a general relationship among species. Relationships between δ ¹⁸O of stem dry matter and stomatal conductance ranged from strongly negative for S. macrophylla to no relationship for T. grandis . Results suggest inter-specific variation among tropical tree species in relationships between stable isotope ratios ( δ ¹³C and δ ¹⁸O) and the gas exchange processes thought to affect them.     

Adaptation to high CO2 concentration in an optimal environment: radiation capture, canopy quantum yield and carbon use efficiency

The effect of elevated [CO₂] on wheat (Triticum aestivum L. Veery 10) productivity was examined by analysing radiation capture, canopy quantum yield, canopy carbon use efficiency, harvest index and daily C gain. Canopies were grown at either 330 or 1200 μ mol mol^–1[CO₂] in controlled environments, where root and shoot C fluxes were monitored continuously from emergence to harvest. A rapidly circulating hydroponic solution supplied nutrients, water and root zone oxygen. At harvest, dry mass predicted from gas exchange data was 102·8 ± 4·7% of the observed dry mass in six trials. Neither radiation capture efficiency nor carbon use efficiency were affected by elevated [CO₂], but yield increased by 13% due to a sustained increase in canopy quantum yield. CO₂ enrichment increased root mass, tiller number and seed mass. Harvest index and chlorophyll concentration were unchanged, but CO₂ enrichment increased average life cycle net photosynthesis (13%, P < 0·05) and root respiration (24%, P < 0·05). These data indicate that plant communities adapt to CO₂ enrichment through changes in C allocation. Elevated [CO₂] increases sink strength in optimal environments, resulting in sustained increases in photosynthetic capacity, canopy quantum yield and daily C gain throughout the life cycle.     

The influence of leaf thickness on the CO2 transfer conductance and leaf stable carbon isotope ratio for some evergreen tree species in Japanese warm-temperate forests

1. The influence of leaf thickness on internal conductance for CO₂ transfer from substomatal cavity to chloroplast stroma ( g _i) and carbon isotope ratio (δ¹³C) of leaf dry matter was investigated for some evergreen tree species from Japanese temperate forests. g _i was estimated based on the combined measurements of gas exchange and concurrent carbon isotope discrimination.
2. Leaves with thicker mesophyll tended to have larger leaf dry mass per area (LMA), larger surface area of mesophyll cells exposed to intercellular air spaces per unit leaf area ( S _mes) and smaller volume ratio of intercellular spaces to the whole mesophyll (mesophyll porosity).
3. g _i of these leaves was correlated positively to S _mes but negatively to mesophyll porosity. The variation in g _i among these species would be therefore primarily determined by variation of the conductance in liquid phase rather than that in gas phase.
4. δ¹³C was positively correlated to mesophyll thickness and leaf nitrogen content on an area basis. However, g _i values did not correlate to δ¹³C. These results suggest that difference in δ¹³C among the species was not caused by the variation in g _i, but mainly by the difference in long-term photosynthetic capacity.
5. Comparison of our results with those of previous studies showed that the correlation between leaf thickness and g _i differed depending on leaf functional types (evergreen, deciduous or annual). Differences in leaf properties among these functional types were discussed.     

In situ studies of water relations and CO2 exchange of the tropical macrolichen, Sticta tomentosa

Diel (24-h) time courses of CO₂ exchange, water relations, and microclimate of the foliose lichen, Sticta tomentosa (Swartz) Ach., and responses to experimentally manipulated conditions were measured at a forest edge in a lower montane rainforest in Panama.
Similar to earlier observations on two other rain forest lichens, daily desiccation suppressed net photosynthesis (NP) during the period when irradiation was highest. Not surprisingly, the light response curves of NP showed saturation at rather low light levels. Rehydration was associated with an initial resaturation burst of short duration, which could be demonstrated both under natural conditions and experimentally. This additional loss of CO₂ seems too low to be ecologically relevant. Moreover, high thallus hydration was also detrimental to NP: at maximum water content net CO₂ uptake was depressed by >50%. Although NP was well adapted to the prevailing high temperatures, the latter also stimulated dark respiration. On average, almost 60% of the diurnal carbon gain was lost during the night.
In spite of these limitations, the integrated 24-h C gain was quite high, on average 0·5% of the thallus C content. Whilst these figures were determined for horizontally exposed samples, we also assessed the role of different exposures on photosynthetic performance. Diel C gain was highest under conditions of semi-shade (westerly exposure), which allowed long periods of activity, whilst much higher irradiance at other exposures could not be utilized for photosynthetic production: easterly exposed thalli dried out even faster than horizontally exposed samples.     

Theoretical reconsiderations when estimating the mesophyll conductance to CO2 diffusion in leaves of C3 plants by analysis of combined gas exchange and chlorophyll fluorescence measurements

Existing methods to estimate the mesophyll conductance to CO₂ diffusion ( g _m) are often based on combined gas exchange and chlorophyll fluorescence measurements. However, estimations of average g _m by these methods are often unreliable either because the range of usable data is too narrow or because the estimations are very sensitive to measurement errors. We describe three method variants to estimate g _m, for which a wider range of data are usable. They use curve-fitting techniques, which minimise the sum of squared model deviations from the data for A (CO₂ assimilation rate) or for J (linear electron transport rate). Like the existing approaches, they are all based on common physiological principles assuming that electron transport limits A . The proposed variants were far less sensitive than the existing approaches to 'measurement noise' either created randomly in the generated data set or inevitably existing in real data sets. Yet, the estimates of g _m from the three variants differed by approximately 15%. Moreover, for each variant, a stoichiometric uncertainty in linear electron transport-limited photosynthesis can cause another 15% difference. Any estimation of g _m using gas exchange and chlorophyll fluorescence measurements should be considered with caution, especially when g _m is high.     

Leaf Gas Exchange Characteristics in Relation to Leaf Canopy Position of Myrica faya in its Native Environment (Tenerife, Canary Islands)

Abstract: Diurnal courses of gas exchange were measured throughout one year in fully expanded current-year leaves in the uppermost canopy (sun leaves, 18 m above ground) and in the lower canopy (shade leaves, 12 m above ground) of Myrica faya Ait., a dominant component of the Canarian laurel forest in Tenerife, Canary Islands, Spain.
M. faya showed large differences between sun and shade leaves in gas exchange characteristics (about 50 % of maximum carbon assimilation rate (A_max) reduction in shade leaves, but this reduction can be higher on specific days) that were modulated by strong light attenuation and high leaf area index (LAI) of the stand. This species presented low A_max, about 10 μmol m^-2 s^-1, high maximum transpiration (E, 8 mmol m^-2 s^-1) and stomatal conductance (g_s, 750 mmol m^-2 s^-1) and very low instantaneous water use efficiency (WUE, mean maximum 1.1 mmol mol^-1) and A/g_s (mean maximum 23.5 μmol mol^-1). M. faya responded to high air vapour pressure deficit (VPD), decreasing its g_s but maintaining relatively high values of A and E during the studied period. Stomatal response to VPD showed a higher sensitivity than its congeners, M. cerifera, and Laurus azorica, tree species co-occurring in the Canarian laurel forest. In general, all these gas exchange characteristics lead us to consider this species more similar to subtropical plants of humid regions than to species of the Mediterranean region.     

Effects of tree height on branch hydraulics, leaf structure and gas exchange in California redwoods

We examined changes in branch hydraulic, leaf structure and gas exchange properties in coast redwood ( Sequoia sempervirens ) and giant sequoia ( Sequoiadendron giganteum ) trees of different sizes. Leaf-specific hydraulic conductivity ( k _L) increased with height in S. sempervirens but not in S. giganteum , while xylem cavitation resistance increased with height in both species. Despite hydraulic adjustments, leaf mass per unit area (LMA) and leaf carbon isotope ratios ( δ ¹³C) increased, and maximum mass-based stomatal conductance ( g _mass) and photosynthesis ( A _mass) decreased with height in both species. As a result, both A _mass and g _mass were negatively correlated with branch hydraulic properties in S. sempervirens and uncorrelated in S. giganteum . In addition, A _mass and g _mass were negatively correlated with LMA in both species, which we attributed to the effects of decreasing leaf internal CO₂ conductance ( g _i). Species-level differences in wood density, LMA and area-based gas exchange capacity constrained other structural and physiological properties, with S. sempervirens exhibiting increased branch water transport efficiency and S. giganteum exhibiting increased leaf-level water-use efficiency with increasing height. Our results reveal different adaptive strategies for the two redwoods that help them compensate for constraints associated with growing taller, and reflect contrasting environmental conditions each species faces in its native habitat.     

Genetic variation in Arabidopsis thaliana for night-time leaf conductance

Night-time leaf conductance ( g _night) and transpiration may have several adaptive benefits related to plant water, nutrient and carbon relations. Little is known, however, about genetic variation in g _night and whether this variation correlates with other gas exchange traits related to water use and/or native habitat climate. We investigated g _night in 12 natural accessions and three near isogenic lines (NILs) of Arabidopsis thaliana . Genetic variation in g _night was found for the natural accessions, and g _night was negatively correlated with native habitat atmospheric vapour pressure deficit (VPD_air), suggesting lower g _night may be favoured by natural selection in drier habitats. However, there were also significant genetic correlations of g _night with daytime gas exchange traits expected to affect plant fitness [i.e. daytime leaf conductance, photosynthesis and intrinsic water-use efficiency (WUE_i)], indicating that selection on daytime gas exchange traits may result in indirect selection on g _night. The comparison of three NILs to their parental genotypes identified one quantitative trait locus (QTL) contributing to variation in g _night. Further characterization of genetic variation in g _night within and among populations and species, and of associations with other traits and native habitats will be needed to understand g _night as a putatively adaptive trait.     

Size-dependent mortality in a Neotropical savanna tree: the role of height-related adjustments in hydraulic architecture and carbon allocation

Size-related changes in hydraulic architecture, carbon allocation and gas exchange of Sclerolobium paniculatum (Leguminosae), a dominant tree species in Neotropical savannas of central Brazil (Cerrado), were investigated to assess their potential role in the dieback of tall individuals. Trees greater than ∼6-m-tall exhibited more branch damage, larger numbers of dead individuals, higher wood density, greater leaf mass per area, lower leaf area to sapwood area ratio (LA/SA), lower stomatal conductance and lower net CO₂ assimilation than small trees. Stem-specific hydraulic conductivity decreased, while leaf-specific hydraulic conductivity remained nearly constant, with increasing tree size because of lower LA/SA in larger trees. Leaves were substantially more vulnerable to embolism than stems. Large trees had lower maximum leaf hydraulic conductance ( K _leaf) than small trees and all tree sizes exhibited lower K _leaf at midday than at dawn. These size-related adjustments in hydraulic architecture and carbon allocation apparently incurred a large physiological cost: large trees received a lower return in carbon gain from their investment in stem and leaf biomass compared with small trees. Additionally, large trees may experience more severe water deficits in dry years due to lower capacity for buffering the effects of hydraulic path-length and soil water deficits.     

Non-stomatal limitation of CO2 assimilation in three tree species during natural drought conditions

The effect of drought on CO₂ assimilation and leaf conductance was studied in three northern hardwood species: Quercus rubra L., Acer rubrum L. and Populus grandidentata Michx. Leaf gas exchange characteristics at two CO₂ levels (320 and 620 μl I⁻¹) and temperatures from 20 to 35°C were measured at the end of a dry period and shortly after 10 cm of rainfall. The effects of drought varied with species, temperature and CO₂ level. Calculated values of internal CO₂ concentration showed little or no decline during drought. Differences in assimilation, before vs after the rains, were most apparent at the higher CO₂ level. These latter two observations indicate nonstomatal disruption of CO₂ assimilation during the dry period. In P. grandidentata there was a substantial interaction between drought and temperature, with a resultant shift in the temperature for maximum assimilation to lower temperatures during drought. During drought, internal CO₂ concentrations increased sharply in all three species under the combined conditions of high temperatures and the higher CO₂ level.     

Short-term effects of aphid feeding on photosynthetic CO2 exchange and dark respiration in legume leaves

Net CO₂ exchange rates and dark respiration rates were determined for single attached legume leaves (leaflets) after 6 to 9 days of aphid infestation. Plant-aphid combinations used were broad bean ( Vicia faba L. cv. Aquadulce) and cowpea [ Vigna unguiculata (L.) Walp. cv. Caloona)] infested with cowpea aphids ( Aphis craccivora Koch) and broad bean and garden pea ( Pisum sativum L. cv. Victory Freezer) infested with pea aphids [ Acyrthosiphon pisum (Harris)]. Leaves from all aphid-infested plants had significantly greater net CO₂ exchange rates in the light than their respective controls and rates of dark respiration of leaves from infested cowpea and garden pea were also significantly greater than those of controls. Dark respiration, as a percentage of net CO₂ exchange rates in the light, was greater in aphid-infested than in control plants. When the mean net daily carbon gain was calculated for the leaves of each plant-aphid combination, leaves from aphid-infested plants had the greatest gain. It is proposed that net CO₂, exchange rates increased due to increased sink demand and dark respiration rates increased to meet the increased energy requirements of phloem loading and cellular maintenance associated with aphid feeding. The apparent compensatory carbon gain of infested leaves was consumed by the aphids.     

Photosynthesis, carbohydrate levels and chlorophyll fluorescence-estimated intercellular CO2 in water-stressed Casuarina equisetifolia Forst. & Forst.

The effect of long-term water stress on photosynthetic carbon metabolism in Casuarina equisetifolia Forst. & Forst. was analysed by measuring CO₂ assimilation, stomatal conductance, the quantum yield of photosystem II ( Φ _PSII), enzyme activities, and the levels of photosynthetic intermediates and carbohydrates. CO₂ assimilation decreased under water stress while the intercellular CO₂ concentration ( C _i) as estimated by gas exchange measurements remained high. However, the estimates of C _i from measurements of Φ _PSII suggest that the decrease in photosynthesis can be explained in terms of stomatal closure. Water stress decreased total stromal fructose-1,6-bisphosphatase activity and did not alter the activities and activation states of ribulose bisphosphate carboxylase oxygenase and NADP-dependent malate dehydrogenase (NADP-MDH). The concentration of photosynthetic metabolites, glucose, fructose and sucrose decreased, whereas starch concentrations increased under drought conditions.     

Variation in growth stimulation by elevated carbon dioxide in seedlings of some C3 crop and weed species

Seven C₃ crop and three C₃ weed species were grown from seed at 360 and at 700 cm³ m^–3 carbon dioxide concentrations in a controlled environment chamber to compare dry mass, relative growth rate (RGR), net assimilation rate (NAR), leaf area ratio (LAR) and photosynthetic acclimation at ambient and elevated carbon dioxide. The dry mass at the final harvest at elevated carbon dioxide relative to that at ambient carbon dioxide was highly correlated with the RGR at the lower carbon dioxide concentration. This relationship could be quite common, because it does not require that species differ in the response of RGR or photosynthesis to elevated carbon dioxide, and holds even when species differ moderately in these responses. RGR was also measured for a limited period at the end of the experiment to determine relationships with leaf gas exchange measured at this time. Relative increases in RGR at elevated carbon dioxide at this time were more highly correlated with the relative increase in NAR at elevated carbon dioxide than with the response of LAR. The amount of acclimation of photosynthesis was a good predictor of the relative increase in NAR at elevated carbon dioxide, and the long-term increase in photosynthesis in the growth environment. No differences between crops and weeds or between cool and warm climate species were found in the responses of growth or photosynthetic acclimation to elevated carbon dioxide.     

Carbon Costs Associated with N2 Fixation in Vicia faba L and Pisum sativum 1. over a 14-Day Period

Abstract: Long-term (14 days) carbon costs of N₂ fixation were studied in pot trials. For this purpose the CO₂ release from the root space of nodulated and non-nodulated (urea nourished) Vicia faba L. and Pisum sativum L. plants was compared and related to the amount of fixed or assimilated N. Additional measurements of shoot CO₂ exchange and dry matter increment were carried out in order to calculate the overall carbon balance. The carbon costs for N₂ fixation in Vicia faba 1. (2.87 mg C/mg N_fiX) were higher than in Pisum sativum L. (2.03 mg C/mg N_fix). However, the better carbon efficiency in Pisum sativum 1. did not lead to a better growth performance compared to Vicia faba L. Vicia faba L. compensated for the carbon and energy expenditure by more intensive photosynthesis in the N₂-fixing treatment. This was not the case with Pisum sativum L., where the carbon balance indicates that the carbon costs of N₂ fixation restricted root growth. It is proposed that low carbon costs for N₂ fixation indicate an adaptation to a critical carbon supply of roots and nodules, e.g., during the pod-filling of grain legumes.