Explore the vast range of titles available.

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, , 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.

Leaf size and venation show remarkable diversity across dicotyledons, and are key determinants of plant adaptation in ecosystems past and present. Here we present global scaling relationships of venation traits with leaf size. Across a new database for 485 globally distributed species, larger leaves had major veins of larger diameter, but lower length per leaf area, whereas minor vein traits were independent of leaf size. These scaling relationships allow estimation of intact leaf size from fragments, to improve hindcasting of past climate and biodiversity from fossil remains. The vein scaling relationships can be explained by a uniquely synthetic model for leaf anatomy and development derived from published data for numerous species. Vein scaling relationships can explain the global biogeographical trend for smaller leaves in drier areas, the greater construction cost of larger leaves and the ability of angiosperms to develop larger and more densely vascularised lamina to outcompete earlier-evolved plant lineages. The size of dicotyledon leaves and their venation vary enormously across ecosystems. In this study, using 485 plant species, scaling relationships are presented between vein traits and leaf size, and explained based on a developmental algorithm that demonstrates why smaller leaves persist in drier areas.

• Hydraulic dysfunction in leaves determines key aspects of whole‐plant responses to water stress; however, our understanding of the physiology of hydraulic dysfunction and its relationships to leaf structure and ecological strategy remains incomplete. • Here, we studied a morphologically and ecologically diverse sample of angiosperms to test whether the water potential inducing a 50% loss in leaf hydraulic conductance (P50leaf) is predicted by properties of leaf xylem relating to water tension‐induced conduit collapse. We also assessed the relationships between P50leaf and other traits considered to reflect drought resistance and ecological strategy. • Across species, P50leaf was strongly correlated with a theoretical predictor of vulnerability to cell collapse in minor veins (the cubed ratio of the conduit wall thickness to the conduit lumen breadth). P50leaf was also correlated with mesophyll traits known to be related to drought resistance, but unrelated to traits associated with carbon economy. • Our data indicate a link between the structural mechanics of leaf xylem and hydraulic function under water stress. Although it is possible that collapse may contribute directly to dysfunction, this relationship may also be a secondary product of vascular economics, suggesting that leaf xylem is dimensioned to avoid wall collapse.

Across plant species, leaves vary enormously in their size and their venation architecture, of which one major function is to replace water lost to transpiration. The leaf hydraulic conductance (K(leaf)) represents the capacity of the transport system to deliver water, allowing stomata to remain open for photosynthesis. Previous studies showed that K(leaf) relates to vein density (vein length per area). Additionally, venation architecture determines the sensitivity of K(leaf) to damage; severing the midrib caused K(leaf) and gas exchange to decline, with lesser impacts in leaves with higher major vein density that provided more numerous water flow pathways around the damaged vein. Because xylem embolism during dehydration also reduces K(leaf), we hypothesized that higher major vein density would also reduce hydraulic vulnerability. Smaller leaves, which generally have higher major vein density, would thus have lower hydraulic vulnerability. Tests using simulations with a spatially explicit model confirmed that smaller leaves with higher major vein density were more tolerant of major vein embolism. Additionally, for 10 species ranging strongly in drought tolerance, hydraulic vulnerability, determined as the leaf water potential at 50% and 80% loss of K(leaf), was lower with greater major vein density and smaller leaf size (vertical bar r vertical bar = 0.85-0.90; P < 0.01). These relationships were independent of other aspects of physiological and morphological drought tolerance. These findings point to a new functional role of venation architecture and small leaf size in drought tolerance, potentially contributing to well-known biogeographic trends in leaf size.

Eudicot leaves have astoundingly diverse shapes. The central problem addressed in this paper is the developmental origin of this diversity. To investigate this problem, we propose a computational model of leaf development that generalizes the largely conserved molecular program for the reference plants Arabidopsis thaliana, Cardamine hirsuta and Solanum lycopersicum. The model characterizes leaf development as a product of three interwoven processes: the patterning of serrations, lobes and/or leaflets on the leaf margin; the patterning of the vascular system; and the growth of the leaf blade spanning the main veins. The veins play a significant morphogenetic role as a local determinant of growth directions. We show that small variations of this model can produce diverse leaf shapes, from simple to lobed to compound. It is thus plausible that diverse shapes of eudicot leaves result from small variations of a common developmental program.

C ₄ photosynthesis is a series of anatomical and biochemical modifications to the typical C ₃ pathway that increases the productivity of plants in warm, sunny, and dry conditions. Despite its complexity, it evolved more than 62 times independently in flowering plants. However, C ₄ origins are absent from most plant lineages and clustered in others, suggesting that some characteristics increase C ₄ evolvability in certain phylogenetic groups. The C ₄ trait has evolved 22–24 times in grasses, and all origins occurred within the PACMAD clade, whereas the similarly sized BEP clade contains only C ₃ taxa. Here, multiple foliar anatomy traits of 157 species from both BEP and PACMAD clades are quantified and analyzed in a phylogenetic framework. Statistical modeling indicates that C ₄ evolvability strongly increases when the proportion of vascular bundle sheath (BS) tissue is higher than 15%, which results from a combination of short distance between BS and large BS cells. A reduction in the distance between BS occurred before the split of the BEP and PACMAD clades, but a decrease in BS cell size later occurred in BEP taxa. Therefore, when environmental changes promoted C ₄ evolution, suitable anatomy was present only in members of the PACMAD clade, explaining the clustering of C ₄ origins in this lineage. These results show that key alterations of foliar anatomy occurring in a C ₃ context and preceding the emergence of the C ₄ syndrome by millions of years facilitated the repeated evolution of one of the most successful physiological innovations in angiosperm history.

The processes by which the functions of interdependent tissues are coordinated as lineages diversify are poorly understood. Here, we examine evolutionary coordination of vascular, epidermal and cortical leaf tissues in the anatomically, ecologically and morphologically diverse woody plant family Proteaceae. We found that, across the phylogenetic range of Proteaceae, the sizes of guard, epidermal, palisade and xylem cells were positively correlated with each other but negatively associated with vein and stomatal densities. The link between venation and stomata resulted in a highly efficient match between potential maximum water loss (determined by stomatal conductance) and the leaf vascular system's capacity to replace that water. This important linkage is likely to be driven by stomatal size, because spatial limits in the packing of stomata onto the leaf surface apparently constrain the maximum size and density of stomata. We conclude that unified evolutionary changes in cell sizes of independent tissues, possibly mediated by changes in genome size, provide a means of substantially modifying leaf function while maintaining important functional links between leaf tissues. Our data also imply the presence of alternative evolutionary strategies involving cellular miniaturization during radiation into closed forest, and cell size increase in open habitats.

High vein density (DV) evolution in angiosperms represented a key functional transition. Yet, a mechanistic account on how this hydraulic transformation evolved remains lacking. We demonstrate that a consequence of producing high DV is that veins must become very small to fit inside the leaf, and that angiosperms are the only clade that evolved the specific type of vessel required to yield sufficiently conductive miniature leaf veins. From 111 species spanning key divergences in vascular plant evolution, we show, using analyses of vein conduit evolution in relation to vein packing, that a key xylem innovation associated with high DV evolution is a strong reduction in vein thickness and simplification of the perforation plates of primary xylem vessels. Simple perforation plates in the leaf xylem occurred only in derived angiosperm clades exhibiting high DV (> 12 mm mm⁻²). Perforation plates in the vessels of other species, including extant basal angiosperms, consisted of resistive scalariform types that were associated with thicker veins and much lower DV. We conclude that a reduction in within‐vein conduit resistance allowed vein size to decrease. We suggest that this adaptation may have been a critical evolutionary step that enabled dramatic DV elaboration in angiosperms.

The leaf economics spectrum (LES) describes large cross-species variation in suites of leaf functional traits ranging from resource-acquisitive to resource-conservative strategies. Such strategies have been integral in explaining plant adaptation to diverse environments, and have been linked to numerous ecosystem processes. The LES has previously been found to be significantly modulated by climate, soil fertility, biogeography, growth form, and life history. One largely unexplored aspect of LES variation, whole-plant ontogeny, is investigated here using multiple populations of three very different species of sunflower: Helianthus annuus, Helianthus mollis, and Helianthus radula. Plants were grown under environmentally controlled conditions and assessed for LES and related traits at four key developmental stages, using recently matured leaves to standardize for leaf age. Nearly every trait exhibited a significant ontogenetic shift in one or more species, with trait patterns differing among populations and species. Photosynthetic rate, leaf nitrogen concentration, and leaf mass per area exhibited surprisingly large changes, spanning over two-thirds of the original cross-species LES variation and shifting from resource-acquisitive to resource-conservative strategies as the plants matured. Other traits being investigated in relation to the LES, such as leaf water content, pH, and vein density, also showed large changes. The finding that ontogenetic variation in LES strategy can be substantial leads to a recommendation of standardization by developmental stage when assessing ‘species values’ of labile traits for comparative approaches. Additionally, the substantial ontogenetic trait shifts seen within single individuals provide an opportunity to uncover the contribution of gene regulatory changes to variation in LES traits.

Leaf veins supply the mesophyll with water that evaporates when stomata are open to allow CO₂ uptake for photosynthesis. Theoretical analyses suggest that water is optimally distributed in the mesophyll when the lateral distance between veins (d x) is equal to the distance from these veins to the epidermis (d y), expressed as d x:d y ≈ 1. Although this theory is supported by observations of many derived angiosperms, we hypothesize that plants in arid environments may reduce d x:d y below unity owing to climate-specific functional adaptations of increased leaf thickness and increased vein density. To test our hypothesis, we assembled leaf hydraulic, morphological, and photosynthetic traits of 68 species from the Eucalyptus and Corymbia genera (termed eucalypts) along an aridity gradient in southwestern Australia. We inferred the potential gas-exchange advantage of reducing d x beyond d y using a model that links leaf morphology and hydraulics to photosynthesis. Our observations reveal that eucalypts in arid environments have thick amphistomatous leaves with high vein densities, resulting in d x:d y ratios that range from 1.6 to 0.15 along the aridity gradient. Our model suggests that, as leaves become thicker, the effect of reducing d x beyond d y is to offset the reduction in leaf gas exchange that would result from maintaining d x:d y at unity. This apparent overinvestment in leaf venation may be explained from the selective pressure of aridity, under which traits associated with long leaf life span, high hydraulic and thermal capacitances, and high potential rates of leaf water transport confer a competitive advantage.