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.
With data from 15 species in eight families of tropical dry forest trees, we provide evidence of coordination between the stem and leaf economic spectra. Species with low-density, flexible, breakable, hydraulically efficient but cavitationally vulnerable wood shed their leaves rapidly in response to drought and had low leaf mass per area and dry mass content. In contrast, species with the opposite xylem syndrome shed their costlier but more drought-resistant leaves late in the dry season. Our results explain variation in the timing of leaf shedding in tropical dry forests: selection eliminates combinations such as low-productivity leaves atop highly vulnerable xylem or water-greedy leaves supplied by xylem of low conductive efficiency. Across biomes, rather than a fundamental trade-off underlying a single axis of trait covariation, the relationship between leaf and stem economics is likely to occupy a wide space in which multiple combinations are possible.
SummaryThe causes underlying bark diversity are unclear. Variation has been frequently attributed to environmental differences across sites. However, variation may also result from tradeoffs and coordination between bark's multiple functions. Bark traits may also covary with wood and leaf traits as part of major dimensions of plant variation.To assess hypotheses regarding tradeoffs and functional coordination, we measured bark traits reflecting protection, storage, mechanics, and photosynthesis in branches of 90 species spanning a wide phylogenetic and environmental range. We also tested associations between bark, wood, and leaf traits. We partitioned trait variation within species, and within and across communities to quantify variation associated with across-site differences.We observed associations between bark mechanics and storage, density and thickness, and thickness and photosynthetic activity. Increasing bark thickness contributed significantly to stiffer stems and greater water storage. Bark density, water content, and mechanics covaried strongly with the equivalent wood traits, and to a lesser degree with leaf size, xylem conductivity, and vessel diameter. Most variation was observed within sites and had low phylogenetic signal.Compared with relatively minor across-site differences, tradeoffs and coordination among functions of bark, leaves, and wood are likely to be major and overlooked factors shaping bark ecology and evolution.
Leaf vein traits are implicated in the determination of gas exchange rates and plant performance. These traits are increasingly considered as causal factors affecting the 'leaf economic spectrum' (LES), which includes the light-saturated rate of photosynthesis, dark respiration, foliar nitrogen concentration, leaf dry mass per area (LMA) and leaf longevity. This article reviews the support for two contrasting hypotheses regarding a key vein trait, vein length per unit leaf area (VLA). Recently, Blonder et al. (2011, 2013) proposed that vein traits, including VLA, can be described as the 'origin' of the LES by structurally determining LMA and leaf thickness, and thereby vein traits would predict LES traits according to specific equations. Careful re-examination of leaf anatomy, published datasets, and a newly compiled global database for diverse species did not support the 'vein origin' hypothesis, and moreover showed that the apparent power of those equations to predict LES traits arose from circularity. This review provides a 'flux trait network' hypothesis for the effects of vein traits on the LES and on plant performance, based on a synthesis of the previous literature. According to this hypothesis, VLA, while virtually independent of LMA, strongly influences hydraulic conductance, and thus stomatal conductance and photosynthetic rate. We also review (i) the specific physiological roles of VLA; (ii) the role of leaf major veins in influencing LES traits; and (iii) the role of VLA in determining photosynthetic rate per leaf dry mass and plant relative growth rate. A clear understanding of leaf vein traits provides a new perspective on plant function independently of the LES and can enhance the ability to explain and predict whole plant performance under dynamic conditions, with applications towards breeding improved crop varieties.
The role of plant polyphenols as defenses against insect herbivores is controversial. We combined correlative field studies across three geographic regions (Northern Mexico, Southern Mexico, and Costa Rica) with induction experiments under controlled conditions to search for candidate compounds that might play a defensive role in the foliage of the tropical oak, Quercus oleoides. We quantified leaf damage caused by four herbivore guilds (chewers, skeletonizers, leaf miners, and gall forming insects) and analyzed the content of 18 polyphenols (including hydrolyzable tannins, flavan-3-ols, and flavonol glycosides) in the same set of leaves using high performance liquid chromatography and mass spectrometry. Foliar damage ranged from two to eight percent per region, and nearly 90% of all the damage was caused by chewing herbivores. Damage due to chewing herbivores was positively correlated with acutissimin B, catechin, and catechin dimer, and damage by mining herbivores was positively correlated with mongolinin A. By contrast, gall presence was negatively correlated with vescalagin and acutissimin B. By using redundancy analysis, we searched for the combinations of polyphenols that were associated to natural herbivory: the combination of mongolinin A and acutissimin B had the highest association to herbivory. In a common garden experiment with oak saplings, artificial damage increased the content of acutissimin B, mongolinin A, and vescalagin, whereas the content of catechin decreased. Specific polyphenols, either individually or in combination, rather than total polyphenols, were associated with standing leaf damage in this tropical oak. Future studies aimed at understanding the ecological role of polyphenols can use similar correlative studies to identify candidate compounds that could be used individually and in biologically meaningful combinations in tests with herbivores and pathogens.
Summary1. The mechanical resistance of leaves has key ecological implications but its basis has not been well understood, particularly at the tissue scale. We tested the hypotheses that leaf mechanical resistance should be a function of tissue density, increasing from the lamina to the midrib, and higher in drought-tolerant than drought-avoiding species. 2. In a common garden study, we quantified nine leaf biomechanical traits, including measurements of material and structural resistance, and in addition 17 morphological traits, at the tissue and whole-leaf scales, for 21 species from three semi-arid communities of California, USA. 3. The mechanical properties of leaves depended strongly on tissue density. Material resistance was significantly greater in the midrib than in the leaf lamina, and tissue resistances were significantly correlated among tissues, lower in deciduous coastal sage species and higher in evergreen drought-tolerant chaparral species. The proportion of the biomass invested in the midrib was lower in species bearing midribs and laminas of high material resistance. 4. Our results support the hypothesis of a hierarchical partitioning of leaf mechanical resistance among leaf tissues reflecting the investment of dry mass. Also, our data indicated a mechanical compensation in leaf design, where leaves with high material resistance and density deploy a relatively minor proportion of support tissue in the midrib. Finally, our results establish a quantitative basis for differences among communities in leaf biomechanics. Our results supported the classical characterization of the mediterranean-climate flora of California according to the dramatic increase in the mean leaf mechanical resistance from species of coastal sage to chaparral, with diverse leaf types in the Mojave Desert species.
The variation in leaf mass per area, leaf nutrients (% carbon, nitrogen and phosphorus), and the allometric relation between tree height and diameter of the black mangrove, Avicennia germinans, were explored in nine mangrove forests in similar environments along a 5° latitudinal gradient in the central region of the Gulf of Mexico, as indicated by a southward increase in temperature and precipitation. There was no correlation between leaf nitrogen or phosphorus content and latitude. Leaf mass per area and leaf carbon content were positively correlated with latitude and negatively correlated with temperature and annual rainfall, whereas asymptotic tree height and maximum diameter showed the opposite trend. Such patterns suggest a trade‐off between leaf traits and tree size which may be constrained by the same environmental factors along a dry‐cold to humid‐warm latitudinal gradient.
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