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A surge of papers have reported low leaf vulnerability to xylem embolism during drought. Here, we focus on the less studied, and more sensitive, outside‐xylem leaf hydraulic responses to multiple internal and external conditions. Studies of 34 species have resolved substantial vulnerability to dehydration of the outside‐xylem pathways, and studies of leaf hydraulic responses to light also implicate dynamic outside‐xylem responses. Detailed experiments suggest these dynamic responses arise at least in part from strong control of radial water movement across the vein bundle sheath. While leaf xylem vulnerability may influence leaf and plant survival during extreme drought, outside‐xylem dynamic responses are important for the control and resilience of water transport and leaf water status for gas exchange and growth.

 

Understanding how environmental adaptations mediate plant and ecosystem responses becomes increasingly important under accelerating global environmental change. Multi-stemmed trees, for example, differ in form and function from single-stemmed trees and may possess physiological advantages that allow for persistence during stressful climatic events such as extended drought. Following the worst drought in Hawaii in a century, we examined patterns of stem abundance and turnover in a Hawaiian lowland dry forest (LDF) and a montane wet forest (MWF) to investigate how multi-stemmed trees might influence site persistence, and how stem abundance and turnover relate to key functional traits. We found stem abundance and multi-stemmed trees to be an important component for climate resilience within the LDF. The LDF had higher relative abundance of multi-stemmed trees, stem abundance, and mean stem abundance compared to a reference MWF. Within the LDF, multi-stemmed trees had higher relative stem abundance (i.e., percent composition of stems to the total number of stems in the LDF) and higher estimated aboveground carbon than single-stemmed trees. Stem abundance varied among species and tree size classes. Stem turnover (i.e., change in stem abundance between five-year censuses) varied among species and tree size classes and species mean stem turnover was correlated with mean species stem abundance per tree. At the plot level, stem abundance per tree is also a predictor of survival, though mortality did not differ between multiple- and single-stemmed trees. Lastly, species with higher mean stem abundance per tree tended to have traits associated with a higher light-saturated photosynthetic rate, suggesting greater productivity in periods with higher water supply. Identifying the traits that allow species and forest communities to persist in dry environments or respond to disturbance is useful for forecasting ecological climate resilience or potential for restoration in tropical dry forests. 

Intra‐specific trait variation (ITV) plays a role in processes at a wide range of scales from organs to ecosystems across climate gradients. Yet, ITV remains rarely quantified for many ecophysiological traits typically assessed for species means, such as pressure volume (PV) curve parameters including osmotic potential at full turgor and modulus of elasticity, which are important in plant water relations. We defined a baseline “reference ITV” (ITV_ref) as the variation among fully exposed, mature sun leaves of replicate individuals of a given species grown in similar, well‐watered conditions, representing the conservative sampling design commonly used for species‐level ecophysiological traits. We hypothesized that PV parameters would show low ITV_refrelative to other leaf morphological traits, and that their intraspecific relationships would be similar to those previously established across species and proposed to arise from biophysical constraints. In a database of novel and published PV curves and additional leaf structural traits for 50 diverse species, we found low ITV_reffor PV parameters relative to other morphological traits, and strong intraspecific relationships among PV traits. Simulation modeling showed that conservative ITV_refenables the use of species‐mean PV parameters for scaling up from spectroscopic measurements of leaf water content to enable sensing of leaf water potential.

 

Abstract Drought decreases water transport capacity of leaves and limits gas exchange, which involves reduced leaf leaf hydraulic conductance (Kleaf) in both the xylem and outside-xylem pathways. Some literature suggests that grapevines are hyper-susceptible to drought-induced xylem embolism. We combined Kleaf and gas exchange measurements, micro-computed tomography of intact leaves, and spatially explicit modeling of the outside-xylem pathways to evaluate the role of vein embolism and Kleaf in the responses of two different grapevine cultivars to drought. Cabernet Sauvignon and Chardonnay exhibited similar vulnerabilities of Kleaf and gs to dehydration, decreasing substantially prior to leaf xylem embolism. Kleaf and gs decreased by 80% for both cultivars by Ψ leaf approximately –0.7 MPa and –1.2 MPa, respectively, while leaf xylem embolism initiated around Ψ leaf = –1.25 MPa in the midribs and little to no embolism was detected in minor veins even under severe dehydration for both cultivars. Modeling results indicated that reduced membrane permeability associated with a Casparian-like band in the leaf vein bundle sheath would explain declines in Kleaf of both cultivars. We conclude that during moderate water stress, changes in the outside-xylem pathways, rather than xylem embolism, are responsible for reduced Kleaf and gs. Understanding this mechanism could help to ensure adequate carbon capture and crop performance under drought. 

Allometric relationships among the dimensions of leaves and their cells hold across diverse eudicotyledons, but have remained untested in the leaves of grasses. We hypothesised that geometric (proportional) allometries of cell sizes across tissues and of leaf dimensions would arise due to the coordination of cell development and that of cell functions such as water, nutrient and energy transport, and that cell sizes across tissues would be associated with light‐saturated photosynthetic rate. We tested predictions across 27 globally distributed C₃and C₄grass species grown in a common garden. We found positive relationships among average cell sizes within and across tissues, and of cell sizes with leaf dimensions. Grass leaf anatomical allometries were similar to those of eudicots, with exceptions consistent with the fewer cell layers and narrower form of grass leaves, and the specialised roles of epidermis and bundle sheath in storage and leaf movement. Across species, mean cell sizes in each tissue were associated with light‐saturated photosynthetic rate per leaf mass, supporting the functional coordination of cell sizes. These findings highlight the generality of evolutionary allometries within the grass lineage and their interlinkage with coordinated development and function.