Sean M. Gleason’s research while affiliated with Colorado State University and other places

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Publications (97)


Conceptual model illustrating a proposed relationship between non‐limited transpiration (LT) (upper panel) and LT crop (lower panel) performance as related to soil water savings and investment in early season growth. Crop and weed development are represented by monocotyledon and dicotyledon plant drawings, respectively. Shallow and deep soil water resources are denoted with full circles, representing full soil profiles, and empty or partially empty circles representing depleted or partially depleted soil profiles. The figure illustrates our three key points: non‐LT crops will exhibit more vigorous early season growth than LT crops, resulting from higher rates of daytime transpiration; vigorous early season growth of non‐LT crops will manifest as increased investment in denser and deeper root systems but poorer development of competing weed plants; and improved growth and competitive ability of non‐LT crop plants, compared to LT crops, will result in improved access to deep soil water and more complete extraction of total soil water resources compared to LT crops.
Published reports citing limited transpiration (LT) theory, LT computer simulations, or LT experiments since the year 2000. Different colored symbols denote different disciplines, and symbol size denotes LT citations received within the discipline.
Relationship between stomatal conductance and net CO2 assimilation in a field‐grown C4 (4‐carbon fixation) species (Sorghum bicolor) grown in Greeley, Colorado. Note the two‐phase relationship between these traits – linear at stomatal conductance values less than c. 0.20 mol m⁻² s⁻¹ (broken blue line), and nonlinear at higher values of stomatal conductance (solid red line). The slope of this relationship represents, at all points along the fitted model (including nonlinear portion), the CO2–H2O exchange rate (when vapor pressure deficit = 1 kPa). We should therefore expect the instantaneous ‘benefit’ of limited transpiration (LT) traits (water saved per unit carbon fixed) to increase in the nonlinear portion, whereas we should expect LT traits to result in a near‐proportional decrease in carbon income and growth in the linear portion of the relationship. Gray symbols denote non‐LT genotypes, whereas green symbols denote LT genotypes. The red portion of the fitted model represents the nonlinear portion of the relationship where we should expect the instantaneous benefit of LT traits to be greatest.
Peer‐reviewed limited transpiration studies published between 1980 and 2023. Note the strong, nearly exponential increase in publication rate after 2010. Each symbol represents the number of publications each year, with the symbol size scaled by citations received as of October 2023.
A 50‐year look‐back on the efficacy of limited transpiration traits: does the evidence support the recent surge in interest?
  • Article
  • Publisher preview available

March 2025

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91 Reads

Sean M. Gleason

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Stephanie K. Polutchko

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Brendan S. Allen

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[...]

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Jared J. Stewart

We examine limited transpiration (LT) traits in crop species, which are claimed to conserve early season water for critical late season growth. Despite there being theoretical support for LT crops, we suggest that there is insufficient empirical evidence to support the general acceptance of this theory. Our criticism focuses on two main points: the undervaluation of early season carbon assimilation and investment over the lifetime of the plant; and the overestimation of soil water savings. We argue that forgoing early season water use, and therefore also future investment in deeper and denser roots (improved resource acquisition), will negatively impact plant performance in many soil and climate contexts. Furthermore, we challenge the assumption that conserved soil water remains available for later use without loss, noting significant losses resulting from evaporation and other sinks. We advocate for a re‐evaluation of LT traits, incorporating a balance of water and carbon dynamics throughout a plant's lifetime. We caution against the adoption of LT traits where they have not been empirically evaluated in the soils and climates of interest to individual research and breeding programs. We propose a more physiologically integrated approach to crop improvement, focusing on water extraction efficiency and strategic carbon investment.

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Xylem embolism refilling revealed in stems of a weedy grass

March 2025

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77 Reads

Proceedings of the National Academy of Sciences

Plant hydraulic dysfunction by embolism formation can impair photosynthesis, growth, and reproduction and, in severe cases, lead to death. Embolism reversal, or “refilling,” is a hypothesized adaptive process in which xylem functionality is rapidly and sustainably restored. This study investigated xylem embolism refilling during recovery from severe drought stress using entirely noninvasive measurements of the same plants. These results were considered in relation to functional traits to address long-standing gaps in understanding the consequences of severe drought stress. Leaf and stem xylem embolism as well as transpiration, photosynthesis, and stem water potential were characterized nondestructively on intact barnyard grass plants during an acute drought event. Plants were rewatered and returned to growth conditions for 10 d, during which time recovery of stem xylem embolism and transpiration were monitored. Leaf xylem embolism and declines in leaf gas exchange occurred mostly between −1.0 MPa and −2.0 MPa, whereas stem xylem embolism occurred mostly between −3.0 MPa and −4.0 MPa. In all measured plants, which included embolism levels up to 88%, stem xylem embolism reversed completely within 24 h after rewatering, and this refilling supported recovery of transpiration and growth after plants were returned to growth conditions. This study provides direct evidence of complete and functional stem xylem refilling. These results present a clear need to elucidate underlying mechanisms and the adaptive significance of this phenomenon as well as its prevalence in nature.


Long‐Term in vivo Observation of Maize Leaf Xylem Embolism, Transpiration and Photosynthesis During Drought and Recovery

February 2025

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12 Reads

Plant water transport is essential to maintain turgor, photosynthesis and growth. Water is transported in a metastable state under large negative pressures, which can result in embolism, that is, the loss of function by the replacement of liquid xylem sap with gas, as a consequence of water stress. To avoid experimental artefacts, we used an optical vulnerability system to quantify embolism occurrence across six fully expanded maize leaves to characterize the sequence of physiological responses (photosynthesis, chlorophyll fluorescence, whole‐plant transpiration and leaf inter‐vein distance) in relation to declining water availability and leaf embolism during severe water stress. Additionally, we characterize the recovery of leaf function in the presence of sustained embolism during a 6‐day recovery period. Embolism formation occurred after other physiological processes were substantially depressed and were irreversible upon rewatering. Recovery of transpiration, net CO 2 assimilation and photosystem II efficiency were aligned with the severity of embolism, whereas these traits returned to near pre‐stress levels in the absence of embolism. A better understanding of the relationships between embolism occurrence and downstream physiological processes during stress and recovery is critical for the improvement of crop productivity and resilience.


Indirect effects of soil amendments on plant traits and the microbiome in post-wildfire forest recovery

February 2025

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39 Reads

Increasing wildfires across the western United States are affecting forest resilience, often leading to novel ecosystems after forest regeneration failures. Following fire, soil organic amendments are commonly applied to reduce erosion and accelerate recovery, but treatments effects on vegetation functional strategies and plant-microbe interactions are poorly understood. Here, we assess the impacts of soil amendments on plant and microbial community composition and plant functional traits 13 years following a severe wildfire in Colorado. Treatments included woody mulch, biochar, and mulch + biochar, which had unique effects on soil nutrients, chemistry, moisture, and microbiomes. We considered three plant functional traits: plant height, specific leaf area (SLA), and leaf dry matter content (LDMC), which had limited direct relationships with treatments. However, random forests and structural equation models revealed non-linear and indirect effects of treatments on soil physical and chemical properties and microbial composition, thereby affecting traits. There was a species-specific response of plant traits to soil N form; Vaccinium scoparium responded more to NH4 + and Oreochrysum parryi responded more to NO3-. Interactions among soil moisture, chemistry (pH and cations), and microbiome composition (especially ectomycorrhizal fungi and nitrifying bacteria and archaea) drove variation in plant traits. These findings show that soil amendment effects on plant traits or microbial composition may not be directly detected using simple statistical approaches, but emerge as indirect effects, revealing the multifaceted importance of soil biotic and abiotic characteristics. This work will benefit future research and restoration practices utilizing a trait-based approach to ecosystem recovery and improving forest resilience.


Surface boundary layer stability and meteorological drivers of temporal microclimate variability in a semiarid grassland

January 2025

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50 Reads

Microclimates are driven by fine-scale weather patterns, play a key role in shaping vegetation responses, and can serve as refuges against climate change. However, little is known about the meteorological factors that drive temporal variability across landscapes. In this study, we deployed a network of air temperature (Tair) and relative humidity (RH) sensors across a small grassland watershed in northern Colorado. We installed sensors at 2 m and 10 m heights to assess stability within the surface boundary layer along with anemometers and radiation sensors. We found significant within-field spatial variability with Tair fluctuating by >15°C and RH by >50% at 15 min intervals. The mean difference in Tair between the highest and lowest points in the watershed was 0.29 °C, corresponding to a near-surface lapse rate of 10 °C km-1, higher than the free-air lapse rate of 6.5 °C km-1. Within-field variability was driven primarily by atmospheric stability, being highest during periods of inversion, with secondary effects of wind speed and solar radiation. These variations resulted in annual variability in biophysical metrics within the field, such as growing degree days and potential evapotranspiration, that exceeded 7%. We found that as mean Tair increased, variability in within-field Tair decreased, indicating that warmer temperatures may reduce microclimate stability, with implications for microrefugia and species' range shifts under warming. This study highlights the temporal dynamics of microclimates which require more thorough representation of surface boundary layer conditions, especially the role of vertical temperature gradients, in shaping landscape-scale ecological, hydrological, and biophysical processes.



Some unique anatomical scaling relationships among genera in the grass subfamily Pooideae

October 2024

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11 Reads

AoB Plants

Members of the grass family Poaceae have adapted to a wide range of habitats and disturbance regimes globally. The cellular structure and arrangements of leaves can help explain how plants survive in different climates, but these traits are rarely measured in grasses. Most studies are focused on individual species or distantly related species within Poaceae. While this focus can reveal broad adaptations, it is also likely to overlook subtle adaptations within more closely-related groups (subfamilies, tribes). This study therefore investigated the scaling relationships between leaf size, Vein Length Area (VLA), and vessel size in five genera within the subfamily Pooideae. The scaling exponent of the relationship between leaf area and VLA was -0.46 (+/- 0.21), which is consistent with previous studies. In Poa and Elymus, however, minor vein number and leaf length were uncorrelated, whereas in Festuca these traits were positively correlated (slope = 0.82 +/- 0.8). These findings suggest there are broad-scale and fine-scale variation in leaf hydraulic traits among grasses. Future studies should consider both narrow and broad phylogenetic gradients.


(a) Leaf hydraulic efficiency (Kmax) and safety (P50, leaf water potential at a loss 50% Kmax) plot for all data. (b) Kmax and P50 relationship after being log‐transformed and fit with a standard major axis (SMA) model. (c) Schematic diagram of an SMA model residual, representing variation in Kmax and P50 orthogonal to the fitted trendline. Asterisks indicate levels of significance based on the SMA model (***, P < 0.001).
Relative predictive capacity and significance of the seven climatic variables on the trade‐off residuals were estimated using a linear mixed regression model. The relative importance of each factor is quantified as the percentage of absolute value of each parameter estimate (standardized regression coefficients) to the sum of absolute values of all parameter estimates. Averaged parameter estimates of the model predictors are shown with their associated 95% confidence intervals. Climate abbreviations and units follow those in Table 1. Asterisks indicate levels of significance (*, P < 0.05; ***, P < 0.001).
Predicted interactive effects between mean annual precipitation (MAP) and mean annual solar radiation (SR) on the trade‐off residuals. The effects of the interaction between MAP and SR were estimated using a multiple linear mixed regression model (MLMM). The predicted surface of residuals arising from the interaction between MAP and SR resembles a twisted plane, because there was a significant interaction between MAP and SR based on MLMM (P < 0.05).
Variation in the trade‐off residuals, leaf hydraulic efficiency (Kmax), and safety (P50) across seven biomes (left) (a–c). (d–f) Corresponding density distributions among biomes. The mean values of mean annual precipitation (MAP; mm) and solar radiation (SR; kJ m⁻² d⁻¹) for different biomes are shown in (a). The full name corresponding to each abbreviated biome are shown. The box plots show quartiles for each trait with extreme values as dots, horizontal lines within boxes represent median values. Different letters indicate significant differences among biomes (one‐way analysis of variance with LSD test, P < 0.05).
Precipitation, solar radiation, and their interaction modify leaf hydraulic efficiency–safety trade‐off across angiosperms at the global scale

October 2024

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487 Reads

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3 Citations

In theory, there is a trade‐off between hydraulic efficiency and safety. However, the strength and direction of this trade‐off at the leaf level are not consistent across studies, and habitat climate may impact this trade‐off. We compiled a leaf hydraulic efficiency and safety dataset for 362 species from 81 sites world‐wide, with 280 paired observations of both traits, and tested whether climate was associated with departure from the proposed trade‐off. The leaf hydraulic efficiency–safety trade‐off was weak (R² = 0.144) at the global scale. Mean annual precipitation and solar radiation (SR) modified the trade‐off. Species from dry and high SR habitats (e.g. desert and tropical savanna) were generally located above the trade‐off line, indicating that these species tended to have higher leaf hydraulic safety and efficiency than species from wet habitats with low SR (e.g. subtropical monsoon forest and montane rainforest), which were located below the trade‐off line. Leaves with high vein density, dry leaf mass per area, and osmotic regulation enhanced safety without compromising hydraulic efficiency. Variation in the hydraulic efficiency–safety trade‐off at the leaf level likely facilitates plant survival in specific habitats and allows for a more nuanced view of leaf hydraulic adaption strategies at the global scale.


A test of limited transpiration traits in sorghum to improve late-season water use, photosynthesis, growth, and grain yield in the high plains of Northern Colorado

June 2024

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40 Reads

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1 Citation

Limited transpiration (LT) traits aim to conserve early-season water to benefit late-season grain development. While theoretical and modeling efforts support LT efficacy, empirical tests directly measuring water loss from leaves and canopies are scarce. This study evaluates the performance of LT genotypes in achieving reduced early-season water use and improved late-season growth and yield in semi-arid Colorado. The research involved near-isogenic lines (NILs) derived from sorghum inbred lines, subjected to different irrigation treatments. Measurements included stomatal conductance, net CO2 assimilation, and photosystem II (PSII) efficiency. Results indicate that LT genotypes did not consistently exhibit lower early-season water use or higher late-season growth compared to non-LT genotypes. Early-season water use was positively correlated with above-ground biomass, challenging the assumption that early-season water conservation can be leveraged for late-season benefits. We question the efficacy of LT traits, highlighting the physiological link between water use and carbon gain, and the potential opportunity costs of reduced early-season growth. We suggests that breeding strategies should focus on enhancing deep soil water access and maximizing carbon gain rather than merely reducing transpiration or shifting water use in arid and semi-arid environments.


Comparative venation costs of monocotyledon and dicotyledon species in the eastern Colorado steppe

May 2024

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13 Reads

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2 Citations

Planta

Leaf vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles did not differ between monocot and dicot species in 21 species from the eastern Colorado steppe. Dicots possessed significantly larger minor vein networks than monocots. Across the tree of life, there is evidence that dendritic vascular transport networks are optimized, balancing maximum speed and integrity of resource delivery with minimal resource investment in transport and infrastructure. Monocot venation, however, is not dendritic, and remains parallel down to the smallest vein orders with no space-filling capillary networks. Given this departure from the “optimized” dendritic network, one would assume that monocots are operating at a significant energetic disadvantage. In this study, we investigate whether monocot venation networks bear significantly greater carbon/construction costs per leaf volume than co-occurring dicots in the same ecosystem, and if so, what physiological or ecological advantage the monocot life form possesses to compensate for this deficit. Given that venation networks could also be optimized for leaf mechanical support or provide herbivory defense, we measured the vascular system of both monocot and dicots at three scales to distinguish between leaf investment in mechanical support (macroscopic vein), total transport and capacitance (vascular bundle), or exclusively water transport (xylem) for both parallel and dendritic venation networks. We observed that vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles was not significantly different between monocot species and dicot species. Dicots, however, possess significantly larger minor vein networks than monocots. The 19 species subjected to gas-exchange measurement in the field displayed a broad range of Amax and but demonstrated no significant relationships with any metric of vascular network size in major or minor vein classes. Given that monocots do not seem to display any leaf hydraulic disadvantage relative to dicots, it remains an important research question why parallel venation (truly parallel, down to the smallest vessels) has not arisen more than once in the history of plant evolution.


Citations (68)


... Over the past two decades, research on functional traits has spanned multiple levels-ranging from physiological to organ-level processes, from individual plants to ecosystems, and from aboveground to belowground interactions -extending into diverse ecological disciplines (Sobral, 2021). The main research areas are as follows: (1) the internal relationships and trade-offs among functional traits (Jin et al., 2024;Zhang et al., 2024); (2) the distribution patterns and variation in traits along environmental gradients (Hulshof et al., 2013;Wieczynski et al., 2019); (3) responses of traits to disturbances such as drought, fire, and grazing (Kerr et al., 2023;Wang et al., 2023); (4) studies on functional diversity (Dıáz and Cabido, 2001); and (5) the mechanisms underlying species coexistence (Mello et al., 2020). The leaf traits of woody plants are closely related to changes in environmental factors such as light and precipitation. ...

Reference:

Changes in the functional traits of Thymus mongolicus along environmental gradients and factors influencing the traits of Northern China
Precipitation, solar radiation, and their interaction modify leaf hydraulic efficiency–safety trade‐off across angiosperms at the global scale

... On the contrary, we view the theory not only as sound within the limits of our understanding of plant-soil-atmosphere interactions but also as having been rarely tested, and in our view, yet to be rigorously confirmed empirically. Further, there are soil, climate, and plant conditions under which LT traits could reduce yield rather than increase it, which should be considered so that this strategy is implemented appropriately (Gleason et al., 2024). Nevertheless, LT literature has received an explosion of interest since around the year 2000, with wide acceptance not just within the plant sciences but across other scientific disciplines as well, for example, biotechnology, molecular biology, ecology, engineering, and atmospheric science (Fig. 2). ...

A test of limited transpiration traits in sorghum to improve late-season water use, photosynthesis, growth, and grain yield in the high plains of Northern Colorado

... Namely, this approach assumes that the variation within trees along the axial axis is consistent across different individuals, an assumption that requires more thorough testing. However, conducting such tests would necessitate the use of additional sensors, which increases research costs or requires expertise to construct them (Drew and Downes 2009, Gleason et al. 2024). This limitation may also reflect the relatively short duration of the study's monitoring period. ...

Development and application of an inexpensive open-source dendrometer for detecting xylem water potential and radial stem growth at high spatial and temporal resolution
  • Citing Article
  • February 2024

AoB Plants

... Moreover, even where studies focused on alteration of root phenotypes, evidence of pot size-induced effects at molecular levels in roots is very limited. Nevertheless, transcriptome-and metabolome-related root traits are now recognised as highly complementary to physiological and biometric ones (Roy and Bassham 2014;Freschet et al. 2021;Drobnitch et al. 2024). The interaction between root growth and pot size is deemed to affect plant cell wall remodelling since roots can sense and respond to mechanical and physical constraints, leading to a reduction in cell elongation and modifications in wall composition (Young et al. 1997;Monshausen and Haswell 2013). ...

Searching for mechanisms driving root pressure in Zea mays—a transcriptomic approach
  • Citing Article
  • March 2024

Journal of Plant Physiology

... In this study, we also found that, for the same phyllotaxy, the diameters of the conduits in the petiolule and main vein, as well as the leaf area, were higher in J. mandshurica than those in the other two species. Previous studies have confirmed that the diameter of the conduit is positively related to leaf area [34][35][36], consistent with our observation, i.e., the leaf area of J. manshurica was higher than the other two species (L8 vs. L5-7 in phyllotaxy). Such findings also supported the classical principle that larger leaves would require a larger 'pipe' system to supply a greater transpiration surface [37]. ...

Grass veins are leaky pipes: vessel widening in grass leaves explain variation in stomatal conductance and vessel diameter among species

... In situations of reduced water availability, trees adopt different strategies to utilize the limited water resources, such as reducing pit size to avoid cavitation (Bueno et al. 2020) and decreasing vessel diameter to prevent embolism and ensure safe transport (Lens et al. 2023). In multi-stemmed coppices, the absorbed water and nutrients are distributed between stems. ...

Comparative anatomy vs mechanistic understanding: how to interpret the diameter-vulnerability link

IAWA Journal

... These regions have experienced increases in the size, duration, and severity of wildfire in recent decades (Calder et al. 2015;Westerling 2016). Though fire is a natural process vital to maintaining ecosystem health in western forests, the changing wildfire regime is likely to impact water quality and availability to western communities (Barnard et al. 2023;Rocca et al. 2014). ...

Wildfire and climate change amplify knowledge gaps linking mountain source-water systems and agricultural water supply in the western United States

Agricultural Water Management

... The changes in these leaf traits represent important photoprotective strategies of plants under drought stress, reducing the risk of photoinhibition or photooxidative stress by limiting light energy input and water loss through evaporation (Bhusal et al., 2021). The adaptation enhances resource conservation and environmental tolerance (Bushey et al., 2023). Additionally, plants growing in drought conditions are generally low in height, which shortens the distance from leaves to rhizosphere moisture, facilitating effective moisture transport (Awad et al., 2010). ...

Water limitation reveals local adaptation and plasticity in the drought tolerance strategies of Bouteloua gracilis

... Recently, the hydraulic conductance of soil and soil-root interface are considered as key factors driving stomatal closure under drought (Cai et al., 2023;Carminati & Javaux, 2020). Consequently, RSM may vary significantly across different soil textures, which deserves further investigation. ...

Soil‐plant hydraulics explain stomatal efficiency‐safety tradeoff

... Based on the assumption that plants will attempt to minimise costs of resource uptake and use based on their specific growing environments, eco-evolutionary optimality principles (Franklin et al. 2020;Harrison et al. 2021;Wright, Reich, and Westoby 2003), which are grounded on optimal coordination (Chen et al. 1993;Maire et al. 2015) and least-cost hypotheses (Wright, Reich, and Westoby 2003), can predict leaf nitrogen demand from climatic variables. Many studies have demonstrated that leaf nitrogen demand for photosynthesis, particularly for ribulose-1,5-biphosphate (RuBP) carboxylase oxygenase (Rubisco) carboxylation (V cmax ), is predominantly determined by climatic factors rather than by soil nitrogen supply (Dong et al. 2017;Onoda et al. 2017;Paillassa et al. 2020;Peng, Bloomfield, and Prentice 2020;Peng et al. 2021;Prentice et al. 2014;Smith et al. 2019;Smith and Keenan 2020;Stocker et al. 2020;Wang et al. 2017;Waring, Perkowski, and Smith 2023;Westerband et al. 2023;Wright, Reich, and Westoby 2003). However, some V cmax variability also appears to be impacted by belowground resources (Paillassa et al. 2020;Smith et al. 2019;Yan et al. 2024). ...

Coordination of photosynthetic traits across soil and climate gradients