Carbon Economy Of Plants Research Articles

Simple and Accurate Representation of Cumulative Nighttime Leaf Respiratory CO2 Efflux.

Leaf respiratory carbon loss decreases independent of temperature as the night progresses. Detailed nighttime measurements needed to quantify cumulative respiratory carbon loss at night are challenging under both lab and field conditions. We provide a simple yet accurate approach to represent variation in nighttime temperature-independent leaf respiratory CO2 efflux in environments with both stable and fluctuating temperatures, which requires no detailed measurements throughout the night. We demonstrate that the inter- and intraspecific variation in the cumulative leaf respiratory CO2 efflux at constant temperature, at any length of night, scales linearly with the inter- and intraspecific variation in initial measurement of leaf respiratory CO2 efflux at the same temperature at the beginning of the night. This approach informs large-scale predictions of cumulative leaf respiratory CO2 efflux, which is needed to understand plant carbon economy in global change studies as well as in global modeling and eddy covariance monitoring of the land-atmosphere exchange of CO2.

Coupled hydraulics and carbon economy underlie age-related growth decline and revitalisation of sand-fixing shrubs after crown removal.

Crown removal revitalises sand-fixing shrubs that show declining vigour with age in drought-prone environments; however, the underlying mechanisms are poorly understood. Here, we addressed this knowledge gap by comparing the growth performance, xylem hydraulics and plant carbon economy across different plant ages (10, 21 and 33 years) and treatments (control and crown removal) using a representative sand-fixing shrub (Caragana microphylla Lam.) in northern China. We found that growth decline with plant age was accompanied by simultaneous decreases in soil moisture, plant hydraulic efficiency and photosynthetic capacity, suggesting that these interconnected changes in plant water relations and carbon economy were responsible for this decline. Following crown removal, quick resprouting, involving remobilisation of root nonstructural carbohydrate reserves, contributed to the reconstruction of an efficient hydraulic system and improved plant carbon status, but this became less effective in older shrubs. These age-dependent effects of carbon economy and hydraulics on plant growth vigour provide a mechanistic explanation for the age-related decline and revitalisation of sand-fixing shrubs. This understanding is crucial for the development of suitable management strategies for shrub plantations constructed with species having the resprouting ability and contributes to the sustainability of ecological restoration projects in water-limited sandy lands.

Trash or treasure: Rhizome conservation during drought

Abstract The role of storage carbohydrates in plant carbon economy is currently disputed as possibly passive accumulation when other resources are limiting growth, or part of a conservative growth strategy as insurance for regrowth and stress response. One indication may be the fate of carbohydrates in senescing rhizomes, as either translocated to be retained in the live and growing end of the rhizome or kept within the senescing rhizome end and lost into the soil for it to decompose. To examine carbohydrate storage in senescing rhizomes, eight rhizomatous species were grown in a split‐pot design with one compartment containing the forward‐growing and younger end of the rhizome and another containing the older end. Both compartments were either watered (control) or the older one was left un‐watered (drought treatment) to trigger rhizome senescence and potential carbohydrate translocation. Plant growth, root traits, and non‐structural carbohydrate types and concentrations were assessed in four sequential harvests. Drought treatment plants had higher rhizome dry matter content. Younger rhizome parts produced higher new rhizome and above‐ground biomass than older rhizome parts. Carbohydrate concentrations in rhizomes remained consistent for both treatments, younger and older rhizome parts, and all harvests, probably because of the translocation of water from the watered to the dry compartment to prevent senescence and rhizome loss. Contrary to expectations, the experimental treatment did not trigger rhizome senescence: plants responded by conserving the rhizome and resources within, rather than by losing their older parts. The invariant composition and concentration of carbohydrates within the rhizome suggest that rhizomes are essential plant organs and the storage carbohydrates they contain are necessary for regrowth after stress. Read the free Plain Language Summary for this article on the Journal blog.

Functional traits predict tree‐level phenological strategies in a mesic Indian savanna

AbstractLeaf phenology influences terrestrial primary production by determining the period of carbon uptake. While dominant leaf habit (evergreen/deciduous) is predictable at large scales based on environmental factors, there is substantial variability in the timing of key events such as leaf flush and senescence at smaller site‐level scales. How much of this variability is explained by species functional traits related to a plant's carbon economy? We monitored leaf phenology for 113 individual trees of eleven dominant species in an Indian mesic savanna. Specifically, we related leaf functional traits and wood density to the (i) timing of leaf flush, leaf maturity, peak canopy, and senescence, (ii) duration of leaf deployment, (iii) duration from start of senescence to leaflessness, and (iv) time to attainment of peak mature canopy following leaf flush. We expected species that use resources conservatively (low specific leaf area (SLA), high leaf dry matter content, low leaf nitrogen, and high wood density) to senesce later and retain their leaves longer. For all species, leaflessness was most pronounced in early dry season and leaf flushing occurred in the late dry season. Species with high leaf carbon and dry matter content showed earlier leaf maturation, attained peak mature canopies sooner, and deployed leaves longer. Species with high SLA, leaf nitrogen, and low wood density showed earlier senescence but relationships were weak. In this savanna, phenology at fine scales was indeed associated with species functional traits relating to carbon investment, but these relationships were strong only when intra‐specific variation in phenology was low.

Non-structural carbohydrate dynamics associated with drought-induced die-off in woody species of a shrubland community.

The relationship between plant carbon economy and drought responses of co-occurring woody species can be assessed by comparing carbohydrate (C) dynamics following drought and rain periods, relating these dynamics to species' functional traits. We studied nine woody species coexisting in a continental Mediterranean shrubland that experienced severe drought effects followed by rain. We measured total non-structural carbohydrates (NSC) and soluble sugars (SS) in roots and stems during drought and after an autumn rain pulse in plants exhibiting leaf loss and in undefoliated ones. We explored whether their dynamics were related to foliage recovery and functional traits (height [H], specific leaf area [SLA], wood density [WD]). During drought, NSC concentrations were overall lower in stems and roots of plants experiencing leaf loss, while SS decreases were smaller. Roots had higher NSC concentrations than stems. After the rain, NSC concentrations continued to decrease, while SS increased. Green foliage recovered after rain, particularly in plants previously experiencing higher leaf loss, independently of NSC concentrations during drought. Species with lower WD tended to have more SS during drought and lower SS increases after rain. In low-WD species, plants with severe leaf loss had lower NSC relative to undefoliated ones. No significant relationship was found between H or SLA and C content or dynamics. Our community-level study reveals that, while responses were species-specific, C stocks overall diminished in plants affected by prolonged drought and did not increase after a pulse of seasonal rain. Dynamics were faster for SS than NSC. We found limited depletion of SS, consistent with their role in basal metabolic, transport and signalling functions. In a scenario of increased drought under climate change, NSC stocks in woody plants are expected to decrease differentially in coexisting species, with potential implications for their adaptive abilities and community dynamics.

Xylem hydraulic safety and construction costs determine tropical tree growth.

Faster growth in tropical trees is usually associated with higher mortality rates, but the mechanisms underlying this relationship are poorly understood. In this study, we investigate how tree growth patterns are linked with environmental conditions and hydraulic traits, by monitoring the cambial growth of 9 tropical cloud forest tree species coupled with numerical simulations using an optimization model. We find that fast-growing trees have lower xylem safety margins than slow-growing trees and this pattern is not necessarily linked to differences in stomatal behaviour or environmental conditions when growth occurs. Instead, fast-growing trees have xylem vessels that are more vulnerable to cavitation and lower density wood. We proposethe growth - xylem vulnerability trade-off represents a wood hydraulic economics spectrum similar to the classic leaf economic spectrum, and show through numerical simulations that this trade-off can emerge from the coordination between growth rates, wood density, and xylem vulnerability to cavitation. Our results suggest that vulnerability to hydraulic failure might be related with the growth-mortality trade-off in tropical trees, determining important life history differences. These findings are important in furthering our understanding of xylem hydraulic functioning and its implications on plant carbon economy.

Resource partitioning by evergreen and deciduous species in a tropical dry forest.

Niche differentiation can lead to coexistence of plant species by partitioning limiting resources. Light partitioning promotes niche differentiation in tropical humid forests, but it is unclear how niche partitioning occurs in tropical dry forests where both light and soil resources can be limiting. We studied the adult niche of four dominant evergreen (cycad, palm) and drought-deciduous (legume, oak) species co-occurring along environmental gradients. We analyzed light intensity and soil fertility effects on key functional traits related to plant carbon and water economy, how these traits determine species' functional strategies, and how these strategies relate to relative species abundance and spatial patterns. Light intensity was negatively associated with a key trait linked to plant water economy (leaf δ 13 C, a proxy for long-term water-use efficiency-WUE), while soil fertility was negatively associated with a key trait for plant carbon economy (LNC, leaf nitrogen content). Evergreens were highly sclerophyllous and displayed an efficient water economy but poor carbon economy, in agreement with a conservative resource-use strategy (i.e., high WUE but low LNC, photosynthetic rates and stature). Conversely, deciduous species, with an efficient carbon economy but poor water economy, exhibited an exploitative resource-use strategy (i.e., high LNC, photosynthetic rates and stature, but low WUE). Evergreen and deciduous species segregated spatially, particularly at fine-scales, as expected for species with different resource-use strategies. The efficient water economy of evergreens was related to their higher relative abundance, suggesting a functional advantage against drought-deciduous species in water-limited environments within seasonally dry tropical forests.

Long-term effect of carbohydrate reserves on growth and reproduction of Prosopis denudans (Fabaceae): implications for conservation of woody perennials.

Prosopis denudans, an extreme xerophyte shrub, is consumed by ungulates and threatened by firewood gathering, because it is one of the preferred species used by Mapuche indigenous people of Patagonia. In a scenario of uncontrolled use of vegetation, it is very difficult to develop a conservation plan that jointly protects natural resources and its users. We performed a field experiment to assess the impact of defoliation on growth, reproduction and stores of a wild population of P. denudans. We imposed four levels of defoliation (removal of 100, 66, 33 and 0% of leaves) and evaluated the short- and long-term (3 years) effects of this disturbance. Seasonal changes in shoot carbohydrates suggested that they support leaf-flush and blooming. Severely defoliated individuals also used root reserves to support growth and leaf-flush after clipping. Vegetative growth was not affected by defoliation history. Leaf mass area increased after the initial clipping, suggesting the development of structural defenses. The depletion of root reserves at the end of the first year affected inflorescence production the following spring. We conclude that P. denudans shrubs could lose up to one-third of their green tissues without affecting growth or inflorescence production. The removal of a higher proportion of leaves will diminish stores, which in turn, will reduce or completely prevent blooming and, therefore, fruit production the following seasons. Very few studies integrate conservation and plant physiology, and we are not aware, so far, of any work dealing with long-term plant carbon economy of a long-lived perennial shrub as an applied tool in conservation. These results might help the development of management strategies that consider both the use and the conservation of wild populations of P. denudans.

How important is woody tissue photosynthesis in poplar during drought stress?

Woody tissue photosynthesis might play a key role in maintaining plant carbon economy and hydraulic function under unfavourable conditions such as drought stress. Within trees, a portion of respired CO2 is assimilated by bark and woody tissue photosynthesis, but its physiological role remains unclear, in particular under unfavour able conditions like drought stress. We hypothesised that woody tissue photosynthesis will contribute to overall tree carbon gain both under sufficient water supply and during drought, and plays a role in maintaining the hydraulic function. We subjected half of the trees to a stem and branch light-exclusion treatment to prevent bark and woody tissue photosynthesis. Then, we measured leaf gas exchange and stem growth in Populus deltoides x nigra ‘Monviso’ trees both under well-watered and dry conditions. We additionally measured cavitation using acoustic emission in detached control and light-excluded branches to illustrate the role of woody tissue photosynthesis in xylem embolism repair. Under well-watered conditions, light exclusion resulted in reduced stem growth relative to control trees by 30 %. In response to drought, stem shrinkage of light-excluded trees was more pronounced as compared to control trees. During drought stress also maximum photosynthesis and transpiration rate tended to decrease more rapidly in light-excluded trees compared to control trees. Leaf fall in light-excluded branches together with the larger number of acoustic emissions in control branches indicates that in the latter more xylem vessels were still hydraulically functional under drought. Therefore, our study highlights that photosynthesis at branch and stem level might be a key factor in the resilience of trees to drought stress by maintaining both the plant carbon economy and hydraulic function.

The enigma of the rise of angiosperms: can we untie the knot?

Multiple hypotheses have been put forward to explain the rise of angiosperms to ecological dominance following the Cretaceous. A unified scheme incorporating all these theories appears to be an inextricable knot of relationships, processes and plant traits. Here, we revisit these hypotheses, categorising them within frameworks based on plant carbon economy, resistance to climatic stresses, nutrient economy, biotic interactions and diversification. We maintain that the enigma remains unresolved partly because our current state of knowledge is a result of the fragmentary nature of palaeodata. This lack of palaeodata limits our ability to draw firm conclusions. Nonetheless, based on consistent results, some inferences may be drawn. Our results indicate that a complex multidriver hypothesis may be more suitable than any single-driver theory. We contend that plant carbon economy and diversification may have played an important role during the early stages of gymnosperms replacement by angiosperms in fertile tropical sites. Plant tolerance to climatic stresses, plant nutrition, biotic interactions and diversification may have played a role in later stages of angiosperm expansion within temperate and harsh environments. The angiosperm knot remains partly tied, but to unravel it entirely will only be feasible if new discoveries are made by scientific communities.

Carbohydrate storage in meadow plants and its depletion after disturbance: do roots and stem-derived organs differ in their roles?

Storage of carbohydrates in organs protected from disturbance is an important adaptation of plants in disturbed habitats. We carried out a field experiment involving 31 herbaceous plant species in two cultural meadows to find out whether roots or belowground stem-derived organs (stem bases, stem tubers and rhizomes) are the main storage organs, to study how reserves accumulate in individual organs in the long term (growing season) and to ascertain whether meadow abandonment affects the distribution of carbohydrate reserves in plants. We also conducted a 22-day pot experiment with four meadow plant species to determine how removal of roots and aboveground parts affects the use of carbohydrates stored in roots and stem-derived organs in the short term. From the long-term perspective of the field experiment, mowing had a positive effect on the concentration of carbohydrate reserves. From the short-term perspective of the pot experiment, however, the effect on concentration and pools of carbohydrates was negative. In the field experiment, carbohydrate concentrations before winter were generally higher than in mid-season, and more often higher in roots than in stem-derived organs. Roots and stem-derived organs of plants in the pot experiment were depleted similarly after both types of disturbance. Our results indicate a need for including both types of belowground plant organs in future studies of the carbon economy of plants from disturbed habitats.

Bark and woody tissue photosynthesis: a means to avoid hypoxia or anoxia in developing stem tissues.

In woody plants, oxygen transport and delivery via the xylem sap are well described, but the contribution of bark and woody tissue photosynthesis to oxygen delivery in stems is poorly understood. Here, we combined stem chlorophyll fluorescence measurements with microsensor quantifications of bark O2 levels and oxygen gas exchange measurements of isolated current-year stem tissues of beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) to investigate how bark and woody tissue photosynthesis impairs the oxygen status of stems. Measurements were made before bud break, when the axial path of oxygen supply via the xylem sap is impeded. At that time, bark O2 levels showed O2 concentrations below the atmospheric concentration, indicating hypoxic conditions or O2 deficiency within the inner bark, but the values were always far away from anoxic. Under illumination bark and woody tissue photosynthesis rapidly increased internal oxygen concentrations compared with plants in the dark, and thereby counteracted against localised hypoxia. The highest photosynthetic activity and oxygen release rates were found in the outermost cortex tissues. By contrast, rates of woody tissue photosynthesis were considerably lower, due to the high light attenuation of the bark and cortex tissues, as well as resistances in radial oxygen diffusion. Therefore, our results confirm that bark and woody tissue photosynthesis not only play a role in plant carbon economy, but may also be important for preventing low oxygen-limitations of respiration in these dense and metabolically active tissues.

Belowground advantages in construction cost facilitate a cryptic plant invasion.

The energetic cost of plant organ construction is a functional trait that is useful for understanding carbon investment during growth (e.g. the resource acquisition vs. tissue longevity tradeoff), as well as in response to global change factors like elevated CO2 and N. Despite the enormous importance of roots and rhizomes in acquiring soil resources and responding to global change, construction costs have been studied almost exclusively in leaves. We sought to determine how construction costs of aboveground and belowground organs differed between native and introduced lineages of a geographically widely dispersed wetland plant species (Phragmites australis) under varying levels of CO2 and N. We grew plants under ambient and elevated atmospheric CO2, as well as under two levels of soil nitrogen. We determined construction costs for leaves, stems, rhizomes and roots, as well as for whole plants. Across all treatment conditions, the introduced lineage of Phragmites had a 4.3 % lower mean rhizome construction cost than the native. Whole-plant construction costs were also smaller for the introduced lineage, with the largest difference in sample means (3.3 %) occurring under ambient conditions. In having lower rhizome and plant-scale construction costs, the introduced lineage can recoup its investment in tissue construction more quickly, enabling it to generate additional biomass with the same energetic investment. Our results suggest that introduced Phragmites has had an advantageous tissue investment strategy under historic CO2 and N levels, which has facilitated key rhizome processes, such as clonal spread. We recommend that construction costs for multiple organ types be included in future studies of plant carbon economy, especially those investigating global change.

Regulation of leaf hydraulics: from molecular to whole plant levels.

The water status of plant leaves is dependent on both stomatal regulation and water supply from the vasculature to inner tissues. The present review addresses the multiple physiological and mechanistic facets of the latter process. Inner leaf tissues contribute to at least a third of the whole resistance to water flow within the plant. Physiological studies indicated that leaf hydraulic conductance (Kleaf) is highly dependent on the anatomy, development and age of the leaf and can vary rapidly in response to physiological or environmental factors such as leaf hydration, light, temperature, or nutrient supply. Differences in venation pattern provide a basis for variations in Kleaf during development and between species. On a short time (hour) scale, the hydraulic resistance of the vessels can be influenced by transpiration-induced cavitations, wall collapses, and changes in xylem sap composition. The extravascular compartment includes all living tissues (xylem parenchyma, bundle sheath, and mesophyll) that transport water from xylem vessels to substomatal chambers. Pharmacological inhibition and reverse genetics studies have shown that this compartment involves water channel proteins called aquaporins (AQPs) that facilitate water transport across cell membranes. In many plant species, AQPs are present in all leaf tissues with a preferential expression in the vascular bundles. The various mechanisms that allow adjustment of Kleaf to specific environmental conditions include transcriptional regulation of AQPs and changes in their abundance, trafficking, and intrinsic activity. Finally, the hydraulics of inner leaf tissues can have a strong impact on the dynamic responses of leaf water potential and stomata, and as a consequence on plant carbon economy and leaf expansion growth. The manipulation of these functions could help optimize the entire plant performance and its adaptation to extreme conditions over short and long time scales.

Molecular and structural analysis of C4-specific PEPC isoform from Pennisetum glaucum plays a role in stress adaptation

Phosphoenolpyruvate carboxylase is an ubiquitous cytosolic enzyme that catalyzes the ß-carboxylation of phosphoenolpyruvate (PEP) and is encoded by multigene family in plants. It plays an important role in carbon economy of plants by assimilating CO2 into organic acids for subsequent C4 or CAM photosynthesis or to perform several anaplerotic roles in non-photosynthetic tissues. In this study, a cDNA clone encoding for PEPC polypeptide possessing signature motifs characteristic to ZmC4PEPC was isolated from Pennisetum glaucum (PgPEPC). Deduced amino acid sequence revealed its predicted secondary structure consisting of forty alpha helices and eight beta strands is well conserved among other PEPC homologs irrespective of variation in their primary amino acid sequences. Predicted PgPEPC quartenary structure is a tetramer consisting of a dimer of dimers, which is globally akin to maize PEPC crystal structure with respect to major chain folding wherein catalytically important amino acid residues of active site geometry are conserved. Recombinant PgPEPC protein expressed in E. coli and purified to homogeneity, possessed in vitro ß-carboxylation activity that is determined using a coupled reaction converting PEP into malate. Tetramer is the most active form, however, it exists in various oligomeric forms depending upon the protein concentration, pH, ionic strength of the media and presence of its substrate or effecters. Recombinant PgPEPC protein confers enhanced growth advantage to E. coli under harsh growth conditions in comparison to their respective controls; suggesting that PgPEPC plays a significant role in stress adaptation.

Can we estimate forest gross primary production from leaf lifespan? A test in a young Fagus crenata forest

It has been well established that leaf longevity is linked to the carbon economy of plants. We used this relationship to predict leaf lifetime carbon gains from leaf lifespan, and estimated the gross primary production (GPP) of a young deciduous forest of Japanese beech (Fagus crenata) located in central Japan. The light-saturated photosynthetic rates of the leaves were measured repeatedly during the growing season. We used the leaf lifespan to calculate the conversion coefficient from the light-saturated photosynthetic rate into the realized leaf lifetime carbon gain under field conditions. The leaf turnover rate was estimated using litter traps. GPP was estimated as the product of lifetime carbon gain per unit of leaf mass, and the annual leaf turnover rate. The GPP of the forest in 2007 was estimated to be <TEX>$1.2{\times}10^3gCm^{-2}y^{-1}$</TEX>, which was within the range of previously reported GPP values of beech forests in Japan, and was close to the GPP of a European beech forest, as estimated by eddy flux measurements.

Convergence towards higher leaf mass per area in dry and nutrient‐poor habitats has different consequences for leaf life span

Summary 1 Leaf life span (LL) and leaf mass per area (LMA) are fundamental traits in the carbon economy of plants, representing the investment required per unit leaf area (LMA) and the duration of the resulting benefit (LL). Species on dry and infertile soils converge towards higher LMA. It has been generally assumed that this allows species from low-resource habitats to achieve longer average leaf life spans, as LMA and LL are often correlated. 2 Leaf life span and LMA were measured for 75 perennial species from eastern Australia. Species were sampled from nutrient-rich and nutrient-poor sites within high and low rainfall regions. LL and LMA were positively correlated across species within each site. In addition, evolutionary divergences in LL and LMA were correlated within each site, indicating that cross-species relationships were not simply driven by differences between higher taxonomic groups. 3 Within a rainfall zone, LL–LMA combinations shifted as expected along common axes of variation such that species on poorer soils had higher LMA and longer LL, but significantly so only at high rainfall. 4 Low rainfall species were expected to have shorter LL at a given LMA or, equally, require higher LMA to achieve a given LL, i.e. shift to a parallel axis of variation, and this was observed on both nutrient-rich and nutrient-poor soils. On average, 30% higher LMA was seemingly required at dry sites to achieve a given LL. Thus, convergence towards higher LMA has different consequences for leaf life span in dry and nutrient-poor habitats. 5 The broad shifts in LL–LMA combinations between site types were also seen when comparing closely related species-pairs (phylogenetically independent contrasts) occurring on nutrient-rich and nutrient-poor soils (within each rainfall zone), and at high- and low-rainfall sites (at each soil nutrient level).

Carbon exchange rates, chlorophyll content, and carbohydrate status of two forest tree species exposed to carbon dioxide enrichment.

Seedlings of yellow-poplar (Liriodendron tulipifera L.) and white oak (Quercus alba L.) were exposed continuously to one of three CO(2) concentrations in open-top chambers under field conditions and evaluated after 24 weeks with respect to carbon exchange rates (CER), chlorophyll (Chl) content, and diurnal carbohydrate status. Increasing the CO(2) concentration from ambient to +150 or +300 microl l(-1) stimulated CER of yellow-poplar and white oak seedlings by 60 and over 35%, respectively, compared to ambient-grown seedlings. The increases in CER were not associated with a significant change in stomatal conductance and occurred despite a reduction in the amounts of Chl and accessory pigments in the leaves of plants grown in CO(2)-enriched air. Total Chl contents of yellow-poplar and white oak seedlings grown at +300 microl l(-1) were reduced by 27 and over 55%, respectively, compared with ambient-grown seedlings. Yellow-poplar and white oak seedlings grown at +300 microl l(-1) contained 72 and 67% more morning starch, respectively, than did ambient-grown plants. In contrast, yellow-poplar and white oak seedlings grown at +300 microl l(-1) contained 17 and 27% less evening sucrose, respectively, than did plants grown at ambient CO(2) concentration. Diurnal starch accumulation and the subsequent depletion of sucrose contributed to a pronounced increase in the starch/sucrose ratio of plants grown in CO(2)-enriched air. All seedlings exhibited a substantial reduction in dark respiration as CO(2) concentration increased, but the significance of this increase to the carbohydrate status and carbon economy of plants grown in CO(2)-enriched air remains unclear.