Temperature-sensitive biochemical 18 O-fractionation and humidity-dependent attenuation factor are needed to predict δ18 O of cellulose from leaf water in a grassland ecosystem.

We explore here our mechanistic understanding of the environmental and physiological processes that determine the oxygen isotope composition of leaf cellulose (δ18 Ocellulose ) in a drought-prone, temperate grassland ecosystem. A new allocation-and-growth model was designed and added to an 18 O-enabled soil-vegetation-atmosphere transfer model (MuSICA) to predict seasonal (April-October) and multi-annual (2007-2012) variation of δ18 Ocellulose and 18 O-enrichment of leaf cellulose (Δ18 Ocellulose ) based on the Barbour-Farquhar model. Modelled δ18 Ocellulose agreed best with observations when integrated over c. 400 growing-degree-days, similar to the average leaf lifespan observed at the site. Over the integration time, air temperature ranged from 7 to 22°C and midday relative humidity from 47 to 73%. Model agreement with observations of δ18 Ocellulose (R2  = 0.57) and Δ18 Ocellulose (R2  = 0.74), and their negative relationship with canopy conductance, was improved significantly when both the biochemical 18 O-fractionation between water and substrate for cellulose synthesis (εbio , range 26-30‰) was temperature-sensitive, as previously reported for aquatic plants and heterotrophically grown wheat seedlings, and the proportion of oxygen in cellulose reflecting leaf water 18 O-enrichment (1 - pex px , range 0.23-0.63) was dependent on air relative humidity, as observed in independent controlled experiments with grasses. Understanding physiological information in δ18 Ocellulose requires quantitative knowledge of climatic effects on pex px and εbio .

Gross and net nitrogen export from leaves of a vegetative C4 grass.

Nitrogen (N) mobilization from mature leaves plays a key role in supplying amino acids to vegetative and reproductive sinks. However, it is unknown if the mobilized N is predominantly sourced by net N-export (a senescence-related process) or other source of N-export from leaves. We used a new approach to partition gross and net N-export from leaf blades at different developmental stages in Cleistogenes squarrosa (a perennial C4 grass). Net N-export was determined as net loss of leaf N with age, while gross N-export was quantified from isotopic mass balances obtained following 24 h-long 15N-labeling with nitrate on 10-12 developmentally distinct (mature and senescing) leaves of individual major tillers. Net N-export was apparent only in older leaves (leaf no. > 7, with leaves numbered basipetally from the tip of the tiller and leaf no. 2 the youngest fully-expanded leaf), while gross N-export was largely independent of leaf age category and was ∼8.4 times greater than the net N-export of a tiller. At whole-tiller level, N import compensated 88 ±â€¯14 (SE) % of gross N-export of all mature blades leading to a net N-export of 0.51 ±â€¯0.07 (SE) µg h-1 tiller-1. N-import was equivalent to 0.09 ±â€¯0.01 (SE) d-1 of total leaf N, similar to reported rates of leaf protein turnover. Gross N-export from all mature blades of a tiller was ∼1.9-times the total demand of the immature tissues of the same (vegetative) tiller. Significant N-export is evident in all mature blades, and is not limited to senescence conditions, implying a much shorter mean residence time of leaf N than that calculated from net N-export. Gross N-export contributes not only to the N demand of the immature tissues of the same tiller but also to N supply of other sinks, such as newly formed tillers. N dynamics at tiller level is integrated with that of the remainder of the shoot, thus highlights the importance of integration of leaf-, tiller-, and plant-scale N dynamics.

Carbon dynamics in aboveground biomass of co-dominant plant species in a temperate grassland ecosystem: same or different?

Understanding the role of individual organisms in whole-ecosystem carbon (C) fluxes is probably the biggest current challenge in C cycle research. Thus, it is unknown whether different plant community members share the same or different residence times in metabolic (τmetab ) and nonmetabolic (i.e. structural) (τnonmetab ) C pools of aboveground biomass and the fraction of fixed C allocated to aboveground nonmetabolic biomass (Anonmetab ). We assessed τmetab , τnonmetab and Anonmetab of co-dominant species from different functional groups (two bunchgrasses, a stoloniferous legume and a rosette dicot) in a temperate grassland community. Continuous, 14-16-d-long (13) C-labeling experiments were performed in September 2006, May 2007 and September 2007. A two-pool compartmental system, with a well-mixed metabolic and a nonmixed nonmetabolic pool, was the simplest biologically meaningful model that fitted the (13) C tracer kinetics in the whole-shoot biomass of all species. In all experimental periods, the species had similar τmetab (5-8 d), whereas τnonmetab ranged from 20 to 58 d (except for one outlier) and Anonmetab from 7 to 45%. Variations in τnonmetab and Anonmetab were not systematically associated with species or experimental periods, but exhibited relationships with leaf life span, particularly in the grasses. Similar pool kinetics of species suggested similar kinetics at the community level.

Fluxes in central carbohydrate metabolism of source leaves in a fructan-storing C3 grass: rapid turnover and futile cycling of sucrose in continuous light under contrasted nitrogen nutrition status.

This work assessed the central carbohydrate metabolism of actively photosynthesizing leaf blades of a C3 grass (Lolium perenne L.). The study used dynamic (13)C labelling of plants growing in continuous light with contrasting supplies of nitrogen ('low N' and 'high N') and mathematical analysis of the tracer data with a four-pool compartmental model to estimate rates of: (i) sucrose synthesis from current assimilation; (ii) sucrose export/use; (iii) sucrose hydrolysis (to glucose and fructose) and resynthesis; and (iv) fructan synthesis and sucrose resynthesis from fructan metabolism. The contents of sucrose, fructan, glucose, and fructose were almost constant in both treatments. Labelling demonstrated that all carbohydrate pools were turned over. This indicated a system in metabolic steady state with equal rates of synthesis and degradation/consumption of the individual pools. Fructan content was enhanced by nitrogen deficiency (55 and 26% of dry mass at low and high N, respectively). Sucrose content was lower in nitrogen-deficient leaves (2.7 versus 6.7%). Glucose and fructose contents were always low (<1.5%). Interconversions between sucrose, glucose, and fructose were rapid (with half-lives of individual pools ranging between 0.3 and 0.8 h). Futile cycling of sucrose through sucrose hydrolysis (67 and 56% of sucrose at low and high N, respectively) and fructan metabolism (19 and 20%, respectively) was substantial but seemed to have no detrimental effect on the relative growth rate and carbon-use efficiency of these plants. The main effect of nitrogen deficiency on carbohydrate metabolism was to increase the half-life of the fructan pool from 27 to 62 h and to effectively double its size.