Scaling relationships and vessel packing in petioles.

PREMISE OF THE STUDY: While tradeoffs among mechanical and conductive functions have been well investigated in woody stems, these tradeoffs are relatively unexplored in petioles, the structural link between stems and laminas. We investigated size-independent scaling relationships between cross-sectional areas of structural and vascular tissues, relationships between tissue areas of xylem and phloem, vessel packing within xylem, and scaling of vascular and structural tissues with lamina traits. METHODS: We examined allometric relationships among petiole tissues and as a function of lamina and petiole size variation on eleven species of Pelargonium. From transverse sections of methacrylate-embedded tissue, we measured the cross-sectional areas of all tissues within the petiole and vessel lumen, and cell wall areas of each vessel. Allometric scaling relationships were analyzed using standardized major axis regressions. KEY RESULTS: Pelargonium petiole vessels were packed as predicted by Sperry's packing rule for woody stems. In contrast to woody stems, there was no evidence of a tradeoff between vessel area and fiber area. Within cross-sections, more xylem was produced than phloem. Among bundles, xylem and phloem scaling relationships varied with bundle position. Except for lamina dry mass and petiole fiber cross-sectional area, petiole and lamina traits were independent. CONCLUSIONS: Petioles share vascular tissue traits with stems despite derivation from leaf primordia. We did not find evidence for a tradeoff between structural and vascular tissues, in part because fibers occur outside the xylem. We propose this separation of conduction and support underlies observed developmental and evolutionary plasticity in petioles.

Seasonal changes in temperate woody plant phloem anatomy and physiology: implications for long-distance transport.

Seasonal changes in climate are accompanied by shifts in carbon allocation and phenological changes in woody angiosperms, the timing of which can have broad implications for species distributions, interactions and ecosystem processes. During critical transitions from autumn to winter and winter to spring, physiological and anatomical changes within the phloem could impose a physical limit on the ability of woody angiosperms to transport carbon and signals. There is a paucity of the literature that addresses tree (floral or foliar) phenology, seasonal phloem anatomy and seasonal phloem physiology together, so our knowledge of how carbon transport could fluctuate seasonally, especially in temperate climates is limited. We review phloem phenology focussing on how sieve element anatomy and phloem sap flow could affect carbon availability throughout the year with a focus on winter. To investigate whether flow is possible in the winter, we construct a simple model of phloem sap flow and investigate how changes to the sap concentration, pressure gradient and sieve plate pores could influence flow during the winter. Our model suggests that phloem transport in some species could occur year-round, even in winter, but current methods for measuring all the parameters surrounding phloem sap flow make it difficult to test this hypothesis. We highlight outstanding questions that remain about phloem functionality in the winter and emphasize the need for new methods to address gaps in our knowledge about phloem function.