Kinetics and relative significance of remobilized and current C and N incorporation in leaf and root growth zones of Lolium perenne after defoliation: assessment by 13C and 15N steady-state labelling

The contribution of pre-defoliation reserves and current assimilates to leaf and root growth was examined in Lolium perenne L. during regrowth after defoliation. Differential steady-state labelling with ¹³C (CO₂ with δ¹³C = -0.0281 and -0.0088) and ¹⁵N (NO₃^? with 1.0 and 0.368 atom percentage, i.e. δ¹⁵N = 1.742 and 0.0052, respectively) was applied for 2 weeks after defoliation. Rapidly growing tissues were isolated, i.e. the basal elongation and maturation zones of the most rapidly expanding leaves and young root tips, with a biomass turnover rate > 1 d^?1. C and N weights of the elongation zone showed a transient decline. The dry matter and C concentration in fresh biomass of leaf growth zones transiently decreased by up to 25% 2 d after defoliation, while the N concentration remained constant. This ‘dilution’ of growth zone C indicates a decreased net influx of carbohydrates relative to growth-related influx of water and N in expanding cells, immediately after defoliation. Recovery of the total C and N weights of the leaf elongation zone coincided with net incorporation of currently absorbed C and N, as shown by the kinetics of δ¹³C and atom percentage ¹⁵N in the growth zones after defoliation. C isotope discrimination (Δ¹³C) in leaf growth zones was about 23‰, 1–2‰ higher than the Δ in root tips. Δ¹⁵N in the leaf and root growth zones was 10±3‰. The leaf elongation zones (at 0–0.03 m from the tiller base) and the distant root tips (about 0.2 m from the base) exhibited similar kinetics of current C and N incorporation. The amount of pre-defoliation C and N in the growth zones, expressed as a fraction of total C and N, decreased from 1.0 to 0.5 at 3 (C) and 5 (N) d after defoliation, and to 0.1 at 5 (C) and 14 (N) d after defoliation. Thus, the dependence of growth zones on current assimilate supply was significant, and stronger for C than for N. The important roles of current assimilates (as compared to pre-defoliation reserves) and ‘dilution’ of dry matter in regrowth after defoliation are discussed in relation to the method of labelling and the functional and morphological heterogeneity of shoot tissues.     

Carbon mobilization in shoot parts and roots of wheat during grain filling: assessment by 13C/12C steady-state labelling, growth analysis and balance sheets of reserves

cv, cultivar
δ, deviation of C isotope composition from a standard
Δ, C isotope discrimination
WSC, water soluble carbohydrates

Steady-state labelling of all post-anthesis photosynthate of wheat was performed to assess the mobilization of pre-anthesis C (C fixed prior to anthesis) in vegetative plant parts during grain filling. Results were compared with estimates obtained by indirect approaches to mobilization of pre-anthesis C: ‘classical’ growth analysis and balance sheets of water soluble carbohydrates (WSC) and protein. Experiments were performed with two spring wheat cultivars grown with differential nitrogen fertilizer supply in 1991 and 1992. The fraction of pre-anthesis C mobilized in above-ground vegetative biomass ranged between 24 and 34% of total C present at anthesis. Treatment effects on mobilization of pre-anthesis C in total above-ground vegetative biomass were closely related (r² = 0·89) to effects on mobilization of WSC-C plus protein-C (estimated as N mobilized × 3·15). On average, 81% of pre-anthesis C mobilization was attributable to the balance of pre-anthesis WSC (48%) and protein (33%) between anthesis and maturity. In roots, WSC and protein mobilization accounted for only 29% of the loss of pre-anthesis C. Notably, mobilization of pre-anthesis C was 1·4–2·6 times larger than the net loss of C from above-ground vegetative biomass between anthesis and maturity. This discrepancy was mainly due to post-anthesis C accumulation in glumes and stem. Post-anthesis C accumulation was related to continued synthesis of structural biomass after anthesis and accounted for a mean 15% of total C contained in above-ground vegetative plant parts at maturity. A close correspondence between net loss of C and mobilization of pre-anthesis C was only apparent in leaf blades and leaf sheaths. Although balance sheets of WSC and protein also underrated the mobilization of pre-anthesis C by ≈ 19%, they gave a much better estimate of pre-anthesis C mobilization than growth analysis.     

Source-sink relations in Lolium perenne L. as reflected by carbohydrate concentrations in leaves and pseudo-stems during regrowth in a free air carbon dioxide enrichment (FACE) experiment*

The effect of an elevated partial pressure of CO₂ (p_CO2) on carbohydrate concentrations in source leaves and pseudo-stems (stubble) of Lolium perenne L. (perennial ryegrass) during regrowth was studied in a regularly defoliated grass sward in the field. The free air carbon dioxide enrichment (FACE) technology enabled natural environmental conditions to be provided. Two levels of nitrogen (N) supply were used to modulate potential plant growth. Carbohydrate concentrations in source leaves were increased at elevated p_CO2, particularly at low N supply. Elevated leaf carbohydrate concentrations were related to an increased structural carbon (C) to N ratio and thus reflected an increased C availability together with a N-dependent sink limitation. Immediately after defoliation, apparent assimilate export rates (differences in the carbohydrate concentrations of young source leaves measured in the evening and on the following morning) showed a greater increase at elevated p_CO2 than at ambient p_CO2; however, replenishment of carbohydrate reserves was not accelerated. Distinct, treatment-dependent carbohydrate concentrations in pseudo-stems suggested an increasing degree of C-sink limitation from the treatment at ambient p_CO2 with high N supply to that at elevated p_CO2 with low N supply. During two growing seasons, no evidence of a substantial change in the response of the carbohydrate source in L. perenne to elevated p_CO2 was found. Our results support the view that the response of L. perenne to elevated p_CO2 is restricted by a C-sink limitation, which is particularly severe at low N supply.