RESUMEN
This study investigates the functional significance of heterophylly in Ginkgo biloba, where leaves borne on short shoots are ontogenetically distinct from those on long shoots. Short shoots are compact, with minimal internodal elongation; their leaves are supplied with water through mature branches. Long shoots extend the canopy and have significant internodal elongation; their expanding leaves receive water from a shoot that is itself maturing. Morphology, stomatal traits, hydraulic architecture, Huber values, water transport efficiency, in situ gas exchange and laboratory-based steady-state hydraulic conductance were examined for each leaf type. Both structure and physiology differed markedly between the two leaf types. Short-shoot leaves were thinner and had higher vein density, lower stomatal pore index, smaller bundle sheath extensions and lower hydraulic conductance than long-shoot leaves. Long shoots had lower xylem area:leaf area ratios than short shoots during leaf expansion, but this ratio was reversed at shoot maturity. Long-shoot leaves had higher rates of photosynthesis, stomatal conductance and transpiration than short-shoot leaves. We propose that structural differences between the two G. biloba leaf types reflect greater hydraulic limitation of long-shoot leaves during expansion. In turn, differences in physiological performance of short- and long-shoot leaves correspond to their distinct ontogeny and architecture.
Asunto(s)
Ginkgo biloba/anatomía & histología , Ginkgo biloba/fisiología , Hojas de la Planta/anatomía & histología , Hojas de la Planta/fisiología , Agua/fisiología , Gases/metabolismo , Ginkgo biloba/citología , Tamaño de los Órganos , Permeabilidad , Hojas de la Planta/citología , Brotes de la Planta/anatomía & histologíaRESUMEN
Although a variety of leaf characteristics appear to be induced by light environment during development, analysis of ontogenetic changes in living broad leaved trees has suggested that a number of other traits also lumped into the classic 'sun' versus 'shade' morphological distinctions, including leaf size, shape, and vein density, are instead controlled largely by local hydraulic environment within the tree canopy. The regularity in how these traits vary with canopy placement suggests a method for addressing a classic paleobotanical quandary: the stature of the source plant - from herb or shrub to canopy tree - is typically unknown for leaf fossils. The study of Ginkgo here complements previous work on Quercus that indicated that leaves throughout the crown are identical in size and venation at the time of bud break and that morphological adaptation to the local microenvironment takes place largely during the expansion phase after the determination of the vascular architecture is complete. Hence, variation in vein density does not reflect differential vein production so much as the distortion of similar vein networks over different final surface areas driven by variation in local hydraulic supply during expansion. Unlike the diffusely growing leaves of the angiosperm, Quercus, the marginally growing leaves of Ginkgo do show some potential for differential vein production, but expansion effects still dominate. The approach suggested here may prove useful for assessing the likelihood that two distinct fossil morphospecies actually represent leaves of the same plant and to gather information concerning canopy structure from disarticulated leaves.
Asunto(s)
Ambiente , Fósiles , Ginkgo biloba/anatomía & histología , Ginkgo biloba/fisiología , Hojas de la Planta/anatomía & histología , Hojas de la Planta/fisiología , Paleontología , ÁrbolesRESUMEN
The electron microprobe allows elemental abundances to be mapped at the microm scale, but until now high resolution mapping of light elements has been challenging. Modifications of electron microprobe procedure permit fine-scale mapping of carbon. When applied to permineralized fossils, this technique allows simultaneous mapping of organic material, major matrix-forming elements, and trace elements with microm-scale resolution. The resulting data make it possible to test taphonomic hypotheses for the formation of anatomically preserved silicified fossils, including the role of trace elements in the initiation of silica precipitation and in the prevention of organic degradation. The technique allows one to understand the localization of preserved organic matter before undertaking destructive chemical analyses and, because it is nondestructive, offers a potentially important tool for astrobiological investigations of samples returned from Mars or other solar system bodies.