Understanding how plants survive drought and cold is increasingly important as plants worldwide experience dieback with drought in moist places and grow taller with warming in cold ones. Crucial in plant climate adaptation are the diameters of water-transporting conduits. Sampling 537 species across climate zones dominated by angiosperms, we find that plant size is unambiguously the main driver of conduit diameter variation. And because taller plants have wider conduits, and wider conduits within species are more vulnerable to conduction-blocking embolisms, taller conspecifics should be more vulnerable than shorter ones, a prediction we confirm with a plantation experiment. As a result, maximum plant size should be short under drought and cold, which cause embolism, or increase if these pressures relax. That conduit diameter and embolism vulnerability are inseparably related to plant size helps explain why factors that interact with conduit diameter, such as drought or warming, are altering plant heights worldwide.
Angiosperm hydraulic performance is crucially affected by the diameters of vessels, the water conducting conduits in the wood. Hydraulic optimality models suggest that vessels should widen predictably from stem tip to base, buffering hydrodynamic resistance accruing as stems, and therefore conductive path, increase in length. Data from 257 species (609 samples) show that vessels widen as predicted with distance from the stem apex across angiosperm orders, habits and habitats. Standardising for stem length, vessels are only slightly wider in warm/moist climates and in lianas, showing that, rather than climate or habit, plant size is by far the main driver of global variation in mean vessel diameter. Terminal twig vessels become wider as plant height increases, while vessel density decreases slightly less than expected tip to base. These patterns lead to testable predictions regarding evolutionary strategies allowing plants to minimise carbon costs per unit leaf area even as height increases.
Flowering plants predominantly conduct water in tubes known as vessels, with vessel diameter playing a crucial role in plant adaptation to climate and reactions to climate change. The importance of vessels makes it essential to understand how and why vessel diameter, plant height, and other ecological factors are interrelated. Although shoot length is by far the main driver of variation in mean vessel diameter across angiosperms, much remains to be understood regarding the factors accounting for the abundant variation around the y‐axis in plots of mean species vessel diameter against shoot length. Here, we explore the potential role of porosity types, wood density, leaf phenology, background imperforate tracheary element type, vasicentric tracheids, vascular tracheids, perforation plate type, and successive cambia in causing variation in the y‐intercept or slope of the mean species vessel‐diameter– and vessel‐density–shoot‐length associations at the shoot tip and base. We detected numerous cases of ecologically significant variation. For example, latewood vessels of ring porous species scale with a lower slope than earlywood, i.e., latewood vessels are relatively narrow in taller plants. This pattern is likely the result of selection favoring freezing‐induced embolism resistance via narrow vessels. Wood density was negatively associated with vessel diameter, with low wood density plants having wider vessels for a given height. Species with scalariform perforation plates scale with a lower shoot base vessel‐diameter–shoot‐length slope, likely reflecting selection against scalariform plates in wide vessels. In other cases, functional groups scaled similarly. For example, species with successive cambia did not differ from those with conventional single cambia in their mean vessel‐diameter–shoot‐length scaling, rejecting our prediction that species with successive cambia should have narrower vessels for a given shoot length. They did, however, have fewer vessels per unit shoot cross‐sectional area than plants of similar heights, likely because vessels have longer functional lifespans (and therefore are fewer) in species with successive cambia. Our methods illustrate how vessel diameter can be studied taking shoot length into account to detect ecologically important variation and construct theory regarding plant adaptation via the hydraulic system that includes plant size as a vital element.
The rhythm of vascular cambial activity was studied in four species of lianas growing in a lowland tropical rainforest of Mexico. Cambial activity was determined by counting the number of layers of cells in the cambial zone. A greater number of layers was assumed to indicate greater cambial activity. In all four species (Machaerium cobanense, M.floribundum, Gouania lupuloides and Trichostigma octandrum) the cambium is active throughout the year. In three of the species (all but T. octandrum) cambial activity was significantly higher in the April-October rainy period than in the November-March dry period. Cambial activity in Trichostigma octandrum was not significantly associated with the wet or dry season.
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