Michael T. Stevens

Resistance and tolerance in Populus tremuloides: genetic variation, costs, and environmental dependency

Plants defend themselves against herbivores via resistance, which reduces damage, and tolerance, which minimizes the negative effects of damage. Theory predicts the existence of tradeoffs between defense and growth, as well as between resistance and tolerance, that could maintain the genetic variation for resistance and tolerance often observed in plant populations. We examined resistance and tolerance among aspen (Populus tremuloides) trees grown under divergent soil nutrient regimes. This common garden experiment revealed substantial genetic variation for resistance and tolerance under both low- and high-nutrient conditions. Costs of resistance exist, particularly under high-nutrient conditions where allocation to resistance chemicals competes directly with growth for limited carbon resources. We found no significant costs of tolerance, however, under either nutrient condition. Despite genetic variation for both resistance and tolerance, we found no evidence for a tradeoff between these two defense traits suggesting that resistance and tolerance are complementary, rather than mutually exclusive, defenses in aspen.

Genetic variation in postfire aspen seedlings in Yellowstone National Park

A rare episode of regeneration of aspen (Populus tremuloides Michx.) by seeds occurred in Yellowstone National Park (YNP), Wyoming, USA, following extensive fires that occurred in 1988. In 1997, we sampled 410 aspen seedlings from 23 local populations distributed widely across YNP to determine how genetic diversity varies with elevation, substrate, plant competition, ungulate browsing, and geographical location. We employed 132 randomly amplified polymorphic DNA (RAPD) markers based on six primers to show genetic relationships within and among the postfire aspen seedling populations. Measures of genetic variation, including estimates of percentage polymorphic loci, expected heterozygosity, and Nei’s FST, indicated that most of the variation occurred within rather than among local populations. There was no indication of geographical differentiation among sampled populations based on hierarchal estimates of Nei’s FST, neighbour‐joining, or correlations between genetic distance and geographical distance. Even genetically distant populations shared nearly 90% of the same markers. Within plots, the amount of genetic variation decreased slightly in response to increased percentage vegetative cover, mean seedling basal diameter, and mean seedling height. Geological substrate, density of lodgepole pine (Pinus contorta var. latifolia Dougl.) seedlings, browsing intensity, and elevation were not significantly related to levels of genetic variation within the seedling plots. These data suggest that genetic variation and geographical structure among seedling populations may occur over time as the transition from seedling‐dominated stands to clone‐dominated stands occurs.

Science Faculty with Education Specialties

Career dynamics for science faculty with interests in education point the way for developing this nascent career specialty.

Investigation of Science Faculty with Education Specialties within the Largest University System in the United States

Efforts to improve science education include university science departments hiring Science Faculty with Education Specialties (SFES), scientists who take on specialized roles in science education within their discipline. Although these positions have existed for decades and may be growing more common, few reports have investigated the SFES approach to improving science education. We present comprehensive data on the SFES in the California State University (CSU) system, the largest university system in the United States. We found that CSU SFES were engaged in three key arenas including K–12 science education, undergraduate science education, and discipline-based science education research. As such, CSU SFES appeared to be well-positioned to have an impact on science education from within science departments. However, there appeared to be a lack of clarity and agreement about the purpose of these SFES positions. In addition, formal training in science education among CSU SFES was limited. Although over 75% of CSU SFES were fulfilled by their teaching, scholarship, and service, our results revealed that almost 40% of CSU SFES were seriously considering leaving their positions. Our data suggest that science departments would likely benefit from explicit discussions about the role of SFES and strategies for supporting their professional activities.

Widespread distribution and unexpected variation among science faculty with education specialties (SFES) across the United States

College and university science departments are increasingly taking an active role in improving science education. Perhaps as a result, a new type of specialized science faculty position within science departments is emerging—referred to here as science faculty with education specialties (SFES)—where individual scientists focus their professional efforts on strengthening undergraduate science education, improving kindergarten-through-12th grade science education, and conducting discipline-based education research. Numerous assertions, assumptions, and questions about SFES exist, yet no national studies have been published. Here, we present findings from a large-scale study of US SFES, who are widespread and increasing in numbers. Contrary to many assumptions, SFES were indeed found across the nation, across science disciplines, and, most notably, across primarily undergraduate, master of science-granting, and PhD-granting institutions. Data also reveal unexpected variations among SFES by institution type. Among respondents, SFES at master of science-granting institutions were almost twice as likely to have formal training in science education compared with other SFES. In addition, SFES at PhD-granting institutions were much more likely to have obtained science education funding. Surprisingly, formal training in science education provided no advantage in obtaining science education funding. Our findings show that the SFES phenomenon is likely more complex and diverse than anticipated, with differences being more evident across institution types than across science disciplines. These findings raise questions about the origins of differences among SFES and are useful to science departments interested in hiring SFES, scientific trainees preparing for SFES careers, and agencies awarding science education funding.

Fire Drives Transcontinental Variation in Tree Birch Defense against Browsing by Snowshoe Hares

Fire has been the dominant disturbance in boreal America since the Pleistocene, resulting in a spatial mosaic in which the most fire occurs in the continental northwest. Spatial variation in snowshoe hare (Lepus americanus) density reflects the fire mosaic. Because fire initiates secondary forest succession, a fire mosaic creates variation in the abundance of early successional plants that snowshoe hares eat in winter, leading to geographic variation in hare density. We hypothesize that fire is the template for a geographic mosaic of natural selection: where fire is greatest and hares are most abundant, hare browsing has most strongly selected juvenile‐phase woody plants for defense. We tested the hypothesis at multiple spatial scales using Alaska birch (Betula neoalaskana) and white birch (Betula papyrifera). We also examined five alternative hypotheses for geographic variation in antibrowsing defense. The fire‐hare‐defense hypothesis was supported at transcontinental, regional, and local scales; alternative hypotheses were rejected. Our results link transcontinental variation in species interactions to an abiotic environmental driver, fire. Intakes of defense toxins by Alaskan hares exceed those by Wisconsin hares, suggesting that the proposed selection mosaic may coincide with a geographic mosaic of coevolution.

Genotypic Differences and Prior Defoliation Affect Re-Growth and Phytochemistry after Coppicing in Populus tremuloides

Although considerable research has explored how tree growth and defense can be influenced by genotype, the biotic environment, and their interaction, little is known about how genotypic differences, prior defoliation, and their interactive effects persist in trees that re-grow after damage that severs their primary stem. To address these issues, we established a common garden consisting of twelve genotypes of potted aspen (Populus tremuloides) trees, and subjected half of the trees to defoliation in two successive years. At the beginning of the third year, all trees were severed at the soil surface (coppiced) and allowed to regenerate for five months. Afterwards, we counted the number of root and stump sprouts produced and measured the basal diameter (d) and height (h) of the tallest ramet in each pot. We collected leaves one and two years after the second defoliation and assessed levels of phenolic glycosides, condensed tannins, and nitrogen. In terms of re-growth, we found that the total number of sprouts produced varied by 3.6-fold among genotypes, and that prior defoliation decreased total sprout production by 24%. The size (d2h) of ramets, however, did not differ significantly among genotypes or defoliation classes. In terms of phytochemistry, we observed genotypic differences in concentrations of all phytochemicals assessed both one and two years after the second defoliation. Two years after defoliation, we observed effects of prior defoliation in a genotype-by-defoliation interaction for condensed tannins. Results from this study demonstrate that genotypic differences and impacts of prior defoliation persist to influence growth and defense traits in trees even after complete removal of above-ground stems, and thus likely influence productivity and plant-herbivore interactions in forests affected by natural disturbances or actively managed through coppicing.

Fostering Change from Within: Influencing Teaching Practices of Departmental Colleagues by Science Faculty with Education Specialties

Globally, calls for the improvement of science education are frequent and fervent. In parallel, the phenomenon of having Science Faculty with Education Specialties (SFES) within science departments appears to have grown in recent decades. In the context of an interview study of a randomized, stratified sample of SFES from across the United States, we discovered that most SFES interviewed (82%) perceived having professional impacts in the realm of improving undergraduate science education, more so than in research in science education or K-12 science education. While SFES reported a rich variety of efforts towards improving undergraduate science education, the most prevalent reported impact by far was influencing the teaching practices of their departmental colleagues. Since college and university science faculty continue to be hired with little to no training in effective science teaching, the seeding of science departments with science education specialists holds promise for fostering change in science education from within biology, chemistry, geoscience, and physics departments.

Root Chemistry in Populus tremuloides: Effects of Soil Nutrients, Defoliation, and Genotype

Although genetic, environmental, and G x E effects on aboveground phytochemistry have been well documented in trembling aspen (Populus tremuloides), little work has focused on the same factors affecting tissues underground. Belowground plant defenses are likely important mediators of root-feeding herbivores that can strongly influence plant fitness. We used a common garden of potted aspen trees to explore the individual and interactive effects of soil nutrient availability, foliar damage, genotype, and their interactions, on concentrations of phytochemicals in aspen roots. Our common garden experiment employed 12 aspen genotypes that were planted into either low- or high-nutrient soil environments. Half of the trees were subjected to defoliation for two successive years, while the others were protected from damage. At the end of the growing season after the second defoliation, we harvested the trees to obtain root samples for which we assessed levels of phenolic glycosides, condensed tannins, nitrogen, and starch. Phenolic glycosides were most affected by genotype, while the other root phytochemicals were most responsive to soil nutrient conditions. The effects of defoliation were observed in interaction with soil nutrient environment and/or genotype. Interestingly, the effect of defoliation on phenolic glycosides was mediated by soil nutrients, whereas the effect of defoliation on condensed tannins was observed in concert with effects of both soil nutrients and genotype. Comparison of data from this study with an earlier, related study revealed that concentrations of phenolic glycosides and condensed tannins are lower in roots than leaves, and less responsive to defoliation. That soil nutrient environment affects root phytochemical concentrations is not unexpected given the intimate association of roots and soil, but the complex interactions between soil nutrients, aboveground damage, and genotype, and their effects on root phytochemistry, are intriguing. Variation in root chemistry could have wide-reaching effects on soil microbial communities, nutrient cycling, and herbivores. Additionally, the response of phytochemicals to damage across organs can link different, spatially separated herbivores as they use different parts of the same plant resource.

Influence of Boulders on Netleaf Hackberry (Celtis reticulata) Growth and Distribution in the Wasatch Foothills

Abstract. In a landscape, abiotic features, such as boulders, influence microhabitats and consequently affect patterns of vegetation. We hypothesized that boulders in the foothills of the Wasatch Mountains east of Provo, Utah, affected the growth patterns of netleaf hackberry (Celtis reticulata) by providing shade on their north faces. To test this hypothesis, we set up 3 transects 6 m wide and up to 50 m long. Along these transects, we measured all hackberries taller than 30 cm (n = 249). We recorded whether the hackberries grew within 0.5 m of a boulder that was at least 0.5 m along one dimension. We found that hackberries at our study site were more likely to be associated with boulders (n = 225; 90.4%) than to be growing alone (n = 24; 10.7%) (x2 = 162.25, df = 1, P < 0.001). For each hackberry associated with a boulder, we took a direction bearing from the center of the boulder to the place where the hackberry was rooted. We found that hackberries associated with boulders were more likely to grow near the south (n = 92; 40.9%) side than near the north (n = 35; 15.6%), west (n = 55; 24.4%), or east sides (n = 43; 19.1%) (x2 = 33.90, df = 3, P < 0.001). These results suggest that boulders influence patterns of hackberry growth and may actually provide thermal radiation that melts snow in the Wasatch foothills, rather than shade protection as we had originally hypothesized.