Negative allometry of leaf xylem conduit diameter and double-wall thickness: implications for implosion safety.

Xylem conduits have lignified walls to resist crushing pressures. The thicker the double-wall (T) relative to its diameter (D), the greater the implosion safety. Having safer conduits may incur higher costs and reduced flow, while having less resistant xylem may lead to catastrophic collapse under drought. Although recent studies have shown that conduit implosion commonly occurs in leaves, little is known about how leaf xylem scales T vs D to trade off safety, flow efficiency, mechanical support, and cost. We measured T and D in > 7000 conduits of 122 species to investigate how T vs D scaling varies across clades, habitats, growth forms, leaf, and vein sizes. As conduits become wider, their double-cell walls become proportionally thinner, resulting in a negative allometry between T and D. That is, narrower conduits, which are usually subjected to more negative pressures, are proportionally safer than wider ones. Higher implosion safety (i.e. higher T/D ratios) was found in asterids, arid habitats, shrubs, small leaves, and minor veins. Despite the strong allometry, implosion safety does not clearly trade off with other measured leaf functions, suggesting that implosion safety at whole-leaf level cannot be easily predicted solely by individual conduits' anatomy.

Leaf venation network architecture coordinates functional trade-offs across vein spatial scales: evidence for multiple alternative designs.

Variation in leaf venation network architecture may reflect trade-offs among multiple functions including efficiency, resilience, support, cost, and resistance to drought and herbivory. However, our knowledge about architecture-function trade-offs is mostly based on studies examining a small number of functional axes, so we still lack a more integrative picture of multidimensional trade-offs. Here, we measured architecture and functional traits on 122 ferns and angiosperms species to describe how trade-offs vary across phylogenetic groups and vein spatial scales (small, medium, and large vein width) and determine whether architecture traits at each scale have independent or integrated effects on each function. We found that generalized architecture-function trade-offs are weak. Architecture strongly predicts leaf support and damage resistance axes but weakly predicts efficiency and resilience axes. Architecture traits at different spatial scales contribute to different functional axes, allowing plants to independently modulate different functions by varying network properties at each scale. This independence of vein architecture traits within and across spatial scales may enable evolution of multiple alternative leaf network designs with similar functioning.

Why are triploid quaking aspen (Populus tremuloides) common?

PREMISE: Quaking aspen is a clonal tree species that has mixed ploidy, often with high relative abundance of both diploids and triploids but no haploids or tetraploids. Triploids typically have low fertility, leaving their occurrence apparently unlikely from an evolutionary perspective, unless they provide a "triploid bridge" to generating higher-fitness tetraploids-which are not observed in this species. This study focused on how triploidy can be maintained in quaking aspen. METHODS: A computational model was used to simulate gamete production, sexual reproduction, asexual reproduction, parent survival, and offspring survival in a population. All parameters were assumed to be cytotype-dependent and environment-independent. Sampling methods were used to identify parameter combinations consistent with observed cytotype frequencies. RESULTS: Many processes and parameter values were sufficient to yield a moderate frequency of triploids, and very few were necessary. The most plausible route involved higher triploid survival at the parent or offspring stage and limited unreduced gamete production by either diploid or triploid parents. Triploid fertility was helpful but not necessary. CONCLUSIONS: The coexistence of diploids and triploids in quaking aspen is statistically likely and promoted by the existence of commonly observed, long-lived triploid clones. However, other mechanisms not captured by the model related to environmental variation could also occur. Further empirical data or more complex but difficult-to-parameterize models are needed to gain further insight.

Navigation between initial and desired community states using shortcuts.

Ecological management problems often involve navigating from an initial to a desired community state. We ask whether navigation without brute-force additions and deletions of species is possible via: adding/deleting a small number of individuals of a species, changing the environment, and waiting. Navigation can yield direct paths (single sequence of actions) or shortcut paths (multiple sequences of actions with lower cost than a direct path). We ask (1) when is non-brute-force navigation possible?; (2) do shortcuts exist and what are their properties?; and (3) what heuristics predict shortcut existence? Using a state diagram framework applied to several empirical datasets, we show that (1) non-brute-force navigation is only possible between some state pairs, (2) shortcuts exist between many state pairs; and (3) changes in abundance and richness are the strongest predictors of shortcut existence, independent of dataset and algorithm choices. State diagrams thus unveil hidden strategies for manipulating species coexistence and efficiently navigating between states.

Plant water use theory should incorporate hypotheses about extreme environments, population ecology, and community ecology.

Plant water use theory has largely been developed within a plant-performance paradigm that conceptualizes water use in terms of value for carbon gain and that sits within a neoclassical economic framework. This theory works very well in many contexts but does not consider other values of water to plants that could impact their fitness. Here, we survey a range of alternative hypotheses for drivers of water use and stomatal regulation. These hypotheses are organized around relevance to extreme environments, population ecology, and community ecology. Most of these hypotheses are not yet empirically tested and some are controversial (e.g. requiring more agency and behavior than is commonly believed possible for plants). Some hypotheses, especially those focused around using water to avoid thermal stress, using water to promote reproduction instead of growth, and using water to hoard it, may be useful to incorporate into theory or to implement in Earth System Models.

Does plant ecosystem thermoregulation occur? An extratropical assessment at different spatial and temporal scales.

To what degree plant ecosystems thermoregulate their canopy temperature (Tc ) is critical to assess ecosystems' metabolisms and resilience with climate change, but remains controversial, with opinions from no to moderate thermoregulation capability. With global datasets of Tc , air temperature (Ta ), and other environmental and biotic variables from FLUXNET and satellites, we tested the 'limited homeothermy' hypothesis (indicated by Tc & Ta regression slope < 1 or Tc < Ta around midday) across global extratropics, including temporal and spatial dimensions. Across daily to weekly and monthly timescales, over 80% of sites/ecosystems have slopes ≥1 or Tc > Ta around midday, rejecting the above hypothesis. For those sites unsupporting the hypothesis, their Tc -Ta difference (ΔT) exhibits considerable seasonality that shows negative, partial correlations with leaf area index, implying a certain degree of thermoregulation capability. Spatially, site-mean ΔT exhibits larger variations than the slope indicator, suggesting ΔT is a more sensitive indicator for detecting thermoregulatory differences across biomes. Furthermore, this large spatial-wide ΔT variation (0-6°C) is primarily explained by environmental variables (38%) and secondarily by biotic factors (15%). These results demonstrate diverse thermoregulation patterns across global extratropics, with most ecosystems negating the 'limited homeothermy' hypothesis, but their thermoregulation still occurs, implying that slope < 1 or Tc < Ta are not necessary conditions for plant thermoregulation.

Climate lags and genetics determine phenology in quaking aspen (Populus tremuloides).

Spatiotemporal patterns of phenology may be affected by mosaics of environmental and genetic variation. Environmental drivers may have temporally lagged impacts, but patterns and mechanisms remain poorly known. We combine multiple genomic, remotely sensed, and physically modeled datasets to determine the spatiotemporal patterns and drivers of canopy phenology in quaking aspen, a widespread clonal dioecious tree species with diploid and triploid cytotypes. We show that over 391 km2 of southwestern Colorado: greenup date, greendown date, and growing season length vary by weeks and differ across sexes, cytotypes, and genotypes; phenology has high phenotypic plasticity and heritabilities of 31-61% (interquartile range); and snowmelt date, soil moisture, and air temperature predict phenology, at temporal lags of up to 3 yr. Our study shows that lagged environmental effects are needed to explain phenological variation and that the effect of cytotype on phenology is obscured by its correlation with topography. Phenological patterns are consistent with responses to multiyear accumulation of carbon deficit or hydraulic damage.

Process-explicit models reveal pathway to extinction for woolly mammoth using pattern-oriented validation.

Pathways to extinction start long before the death of the last individual. However, causes of early stage population declines and the susceptibility of small residual populations to extirpation are typically studied in isolation. Using validated process-explicit models, we disentangle the ecological mechanisms and threats that were integral in the initial decline and later extinction of the woolly mammoth. We show that reconciling ancient DNA data on woolly mammoth population decline with fossil evidence of location and timing of extinction requires process-explicit models with specific demographic and niche constraints, and a constrained synergy of climatic change and human impacts. Validated models needed humans to hasten climate-driven population declines by many millennia, and to allow woolly mammoths to persist in mainland Arctic refugia until the mid-Holocene. Our results show that the role of humans in the extinction dynamics of woolly mammoth began well before the Holocene, exerting lasting effects on the spatial pattern and timing of its range-wide extinction.

Gas exchange analysers exhibit large measurement error driven by internal thermal gradients.

Portable gas exchange analysers provide critical data for understanding plant-atmosphere carbon and water fluxes, and for parameterising Earth system models that forecast climate change effects and feedbacks. We characterised temperature measurement errors in the Li-Cor LI-6400XT and LI-6800, and estimated downstream errors in derived quantities, including stomatal conductance (gsw ) and leaf intercellular CO2 concentration (Ci ). The LI-6400XT exhibited air temperature errors (differences between reported air temperature and air temperature measured near the leaf) up to 7.2°C, leaf temperature errors up to 5.3°C, and relative errors in gsw and Ci that increased as temperatures departed from ambient. This caused errors in leaf-to-air temperature relationships, assimilation-temperature curves and CO2 response curves. Temperature dependencies of maximum Rubisco carboxylation rate (Vcmax ) and maximum RuBP regeneration rate (Jmax ) showed errors of 12% and 35%, respectively. These errors are likely to be idiosyncratic and may differ among machines and environmental conditions. The LI-6800 exhibited much smaller errors. Earth system model predictions may be erroneous, as much of their parametrisation data were measured on the LI-6400XT system, depending on the methods used. We make recommendations for minimising errors and correcting data in the LI-6400XT. We also recommend transitioning to the LI-6800 for future data collection.

Remote sensing of cytotype and its consequences for canopy damage in quaking aspen.

Mapping geographic mosaics of genetic variation and their consequences via genotype x environment interactions at large extents and high resolution has been limited by the scalability of DNA sequencing. Here, we address this challenge for cytotype (chromosome copy number) variation in quaking aspen, a drought-impacted foundation tree species. We integrate airborne imaging spectroscopy data with ground-based DNA sequencing data and canopy damage data in 391 km2 of southwestern Colorado. We show that (1) aspen cover and cytotype can be remotely sensed at 1 m spatial resolution, (2) the geographic mosaic of cytotypes is heterogeneous and interdigitated, (3) triploids have higher leaf nitrogen, canopy water content, and carbon isotope shifts (δ13 C) than diploids, and (4) canopy damage varies among cytotypes and depends on interactions with topography, canopy height, and trait variables. Triploids are at higher risk in hotter and drier conditions.