The Widened Pipe Model of plant hydraulic evolution.

Shaping global water and carbon cycles, plants lift water from roots to leaves through xylem conduits. The importance of xylem water conduction makes it crucial to understand how natural selection deploys conduit diameters within and across plants. Wider conduits transport more water but are likely more vulnerable to conduction-blocking gas embolisms and cost more for a plant to build, a tension necessarily shaping xylem conduit diameters along plant stems. We build on this expectation to present the Widened Pipe Model (WPM) of plant hydraulic evolution, testing it against a global dataset. The WPM predicts that xylem conduits should be narrowest at the stem tips, widening quickly before plateauing toward the stem base. This universal profile emerges from Pareto modeling of a trade-off between just two competing vectors of natural selection: one favoring rapid widening of conduits tip to base, minimizing hydraulic resistance, and another favoring slow widening of conduits, minimizing carbon cost and embolism risk. Our data spanning terrestrial plant orders, life forms, habitats, and sizes conform closely to WPM predictions. The WPM highlights carbon economy as a powerful vector of natural selection shaping plant function. It further implies that factors that cause resistance in plant conductive systems, such as conduit pit membrane resistance, should scale in exact harmony with tip-to-base conduit widening. Furthermore, the WPM implies that alterations in the environments of individual plants should lead to changes in plant height, for example, shedding terminal branches and resprouting at lower height under drier climates, thus achieving narrower and potentially more embolism-resistant conduits.

The vessel wall thickness-vessel diameter relationship across woody angiosperms.

PREMISE: Comparative anatomy is necessary to identify the extremes of combinations of functionally relevant structural traits, to ensure that physiological data cover xylem anatomical diversity adequately, and thus achieve a global understanding of xylem structure-function relations. A key trait relationship is that between xylem vessel diameter and wall thickness of both the single vessel and the double vessel+adjacent imperforate tracheary element (ITE). METHODS: We compiled a comparative data set with 1093 samples, 858 species, 350 genera, 86 families, and 33 orders. We used broken linear regression and an algorithm to explore changes in parameter values from linear regressions using subsets of the data set to identify a threshold, at 90-µm vessel diameter, in the wall thickness-diameter relationship. RESULTS: Below 90 µm diameter for vessels, virtually any wall thickness could be associated with virtually any diameter. Below this threshold, selection is free to favor a very wide array of combinations, such as very thick walls and narrow vessels in ITE-free herbs, or very thin-walled, wide vessels in evergreen dryland pioneers. Above 90 µm, there was a moderate positive relationship. CONCLUSIONS: Our analysis shows that the space of vessel wall thickness-diameter combinations is very wide, with selection apparently eliminating individuals with vessel walls "too thin" for their diameter. Most importantly, our survey revealed poorly studied plant hydraulic syndromes (functionally significant trait combinations). These data suggest that the full span of trait combinations, and thus the minimal set of hydraulic syndromes requiring study to span woody plant functional diversity adequately, remains to be documented.

ULTRAPETALA1 maintains Arabidopsis root stem cell niche independently of ARABIDOPSIS TRITHORAX1.

During plant development, morphogenetic processes rely on the activity of meristems. Meristem homeostasis depends on a complex regulatory network constituted by different factors and hormone signaling that regulate gene expression to coordinate the correct balance between cell proliferation and differentiation. ULTRAPETALA1, a transcriptional regulatory protein described as an Arabidopsis Trithorax group factor, has been characterized as a regulator of the shoot and floral meristems activity. Here, we highlight the role of ULTRAPETALA1 in root stem cell niche maintenance. We found that ULTRAPETALA1 is required to regulate both the quiescent center cell division rate and auxin signaling at the root tip. Furthermore, ULTRAPETALA1 regulates columella stem cell differentiation. These roles are independent of the ARABIDOPSIS TRITHORAX1, suggesting a different mechanism by which ULTRAPETALA1 can act in the root apical meristem of Arabidopsis. This work introduces a new component of the regulatory network needed for the root stem cell niche maintenance.

The evolution of ontogenetic "decision-making" in the wood of a clade of tropical plants.

Greater diversity in functional morphology should be associated with the evolution of greater ontogenetic diversity, an expectation difficult to test in most long-lived wild organisms. In the cells derived from the wood meristem (vascular cambium), plants provide extraordinary systems for reconstructing ontogenies in often long-lived organisms. The vascular cambium produces files of cells from the stem center to the periphery, with each cambial derivative "deciding" which of four cell types it differentiates into. Wood cell files remain in place, allowing tracing of the ontogenetic "decisions" taken throughout the life of a stem. We compared cell files from the Pedilanthus clade (genus Euphorbia), which span a range of growth forms from small trees and shrubs of tropical habitats to desert succulents. Using language theory, we represented wood cell types as "letters" and combinations of cell types in cell files as "words," allowing us to measure the diversity of decisions based on word frequency matrices. We also used information content metrics to compare levels of predictability in "decision-making." Our analyses identified a wider array of developmental decisions in woody trees as compared to succulent shrubs, illustrating ways that woody plants provide unparalleled systems for studying the evolution of ontogeny in long-lived, non-model species.

Plant morpho evo-devo.

Evo-devo is often thought of as being the study of which genes underlie which phenotypes. However, evo-devo is much more than this, especially in plant science. In leaf scars along stems, cell changes across wood growth rings, or flowers along inflorescences, plants trace a record of their own development. Plant morpho evo-devo provides data that genes could never furnish on themes such as heterochrony, the evolution of temporal phenotypes, modularity, and phenotype-first evolution. As plant science surges into increasingly -omic realms, it is essential to keep plant morpho evo-devo in full view as an honored member of the evo-devo canon, ensuring that plant scientists can, wherever they are, generate fundamental insights at the appropriate level of biological organization.