Leaf transition from heterotrophy to autotrophy is recorded in the intraleaf C, H and O isotope patterns of leaf organic matter

摘要

Rationale Quantitatively relating(13)C/C-12,H-2/H-1 and(18)O/O-16 ratios of plant alpha-cellulose and(2)H/H-1 ofn-alkanes to environmental conditions and metabolic status should ideally be based on the leaf, the plant organ most sensitive to environmental change. The fact that leaf organic matter is composed of isotopically different heterotrophic and autotrophic components means that it is imperative that one be able to disentangle the relative heterotrophic and autotrophic contributions to leaf organic matter. Methods We tackled this issue by two-dimensional sampling of leaf water and alpha-cellulose, and specificn-alkanes from greenhouse-grown immature and mature and field-grown mature banana leaves, taking advantage of their large areas and thick waxy layers. Leaf water, alpha-cellulose andn-alkane isotope ratios were then characterized using elemental analysis isotope ratio mass spectrometry (IRMS) or gas chromatography IRMS. A three-member (heterotrophy, autotrophy and photoheterotrophy) conceptual linear mixing model was then proposed for disentangling the relative contributions of the three trophic modes. Results We discovered distinct spatial leaf water, alpha-cellulose andn-alkane isotope ratio patterns that varied with leaf developmental stages. We inferred from the conceptual model that, averaged over the leaf blade, only 20% of alpha-cellulose in banana leaf is autotrophically laid down in both greenhouse-grown and field-grown banana leaves, while approximately 60% and 100% ofn-alkanes are produced autotrophically in greenhouse-grown and field-grown banana leaves, respectively. There exist distinct lateral (edge to midrib) gradients in autotrophic contributions of alpha-cellulose andn-alkanes. Conclusions Efforts to establish quantitative isotope-environment relationships should take into account the fact that the evaporative leaf water(18)O and(2)H enrichment signal recorded in autotrophically laid down alpha-cellulose is significantly diluted by the heterotrophically formed alpha-cellulose. The delta H-2 value of field-grown mature banana leafn-alkanes is much more sensitive than alpha-cellulose as a recorder of the growth environment. Quantitative isotope-environment relationship based on greenhouse-grownn-alkane delta H-2 values may not be reliable.