Herbivory by wild and domestic ungulates is a chronic disturbance that can have dramatic effects on vegetation dynamics. Herbivory effects, however, are not easily predicted under different combinations of episodic disturbance such as fire, timber harvest, drought, and insect defoliation. This lack of predictability poses a substantial obstacle to effective management of ungulate herbivory. Traditional models of vegetation transition in forested ecosystems have ignored the influences of ungulate herbivory, while research on effects of herbivory have typically excluded other disturbances. Of the 82 contemporary studies on ungulate herbivory we examined, only 15 (18%) considered the interactions of herbivory with episodic disturbances. Moreover, only 26 (32%) evaluated vegetation response to ungulate herbivory beyond the simplistic treatment levels of herbivory versus no herbivory. Only 31 (38%) used a repeated‐measures design of sampling responses over 3 or more time periods. Finally, just 7 (9%) explicitly made inferences to large landscapes such as watersheds, which are often used for management planning. We contend that useful landscape research on herbivory must examine the interactions of ungulate grazing with other disturbance regimes at spatial extents of interest to forest and rangeland managers and under varying ungulate densities and species. We identify herbivory models that could accommodate such information for forested landscapes in western North America. Such models are essential for identifying knowledge gaps, designing future studies, and validating relations of ungulate herbivory on landscapes where episodic disturbances are common, such as those of western North America.

Protein C inhibitor (PCI) inhibits multiple plasma serine proteases. To determine which residues contribute to its specificity of inhibition, 19 mutations in the reactive site loop of PCI (from Thr352 to Arg357) were generated and assayed with thrombin, activated protein C (APC), and factor Xa. To identify the intermolecular interactions responsible for these kinetics, a molecular model of PCI was generated using alpha 1-protease inhibitor and ovalbumin as templates. This model of PCI was docked with thrombin, followed by extensive energy minimization, to determine a lowest energy complex. The resulting docked complex was used as a template to form molecular models of PCI-APC and PCI-factor Xa complexes. The best inhibitors of thrombin contained Pro or Gly at the P2 position in place of Phe353, with 2- and 7-fold increases in activity, respectively. These substitutions reduced steric interactions with the 60-insertion loop unique to thrombin. The best inhibitors of APC and factor Xa contained Arg at the P3 position in place of Thr352, with 2- and 5-fold increases in inhibition rates, respectively. The molecular model predicts that Arg in this position could form a salt bridge with Glu217 of each protease. Changing Arg357 at the P3' position had little effect on protease inhibition, consistent with the observation in the model that this residue points toward the body of PCI, forming a salt bridge with Glu220. Given its broad specificity of inhibition, PCI has proven very useful in understanding the nature of serpin-protease interactions using multiple mutations in a serpin assayed with multiple proteases.

a b s t r a c tHerbivory by domestic and wild ungulates can dramatically affect vegetation structure, composition and dynamics in nearly every terrestrial ecosystem of the world. These effects are of particular concern in forests of western North America, where intensive herbivory by native and domestic ungulates has the potential to substantially reduce or eliminate deciduous, highly palatable species of aspen (Populus tremuloides), cottonwood (Populus trichocarpa), and willow (Salix spp.). In turn, differential herbivory pressure may favor greater establishment of unpalatable conifers that serve as ladder fuels for stand-replacing fires. The resulting high fuel loads often require silvicultural fuels reductions to mitigate fire risk, which in turn may facilitate additional recruitment of deciduous species but also additional herbivory pressure. Potential interactions of ungulate herbivory with episodic disturbances of silviculture, fire, and other land uses are not well documented, but are thought to operate synergistically to affect forest dynamics. We evaluated individual and joint effects of ungulate herbivory and fuels reduction treatments in grand fir (Abies grandis) and Douglas-fir (Psuedotsuga menziezii) forests that dominate large areas of interior western North America. We applied fuels reduction treatments of mechanical thinning and prescribed fire and then evaluated the responses of aspen, cottonwood, and willow species to these treatments (N = 3) versus areas of no treatment (N = 3), and to exclusion from ungulate herbivory versus areas subjected to extant herbivory by free-ranging cattle (Bos taurus), elk (Cervus elaphus), and mule deer (Odocoileus hemionus). Densities of deciduous species were >4 times higher in response to fuels reduction treatments (84.4 individuals/ha) compared to areas of no treatment (19.7 individuals/ha). Additionally, when ungulates were excluded from fuels treated sites, the density of cottonwood was >5 times higher (122.5 individuals/ha) than fuels treated sites subjected to extant herbivory (24.3 individuals/ha). Similarly, densities of Populus spp. and Salix spp. were >3 times higher (211.6 individuals/ha) on fuels treated sites excluded from ungulate herbivory versus fuels treated sites subjected to extant herbivory (66.1 stems/ha). Deciduous species subjected to extant ungulate herbivory also were significantly lower in height, canopy surface area, and canopy volume than the same species inside the ungulate exclosures. Recruitment and long-term survival of aspen, cottonwood, and willow species in coniferous forests of interior western North America require a combination of episodic disturbances such as silviculture and fire to facilitate deciduous plant recruitment, followed by reductions in grazing pressure by domestic and wild ungulates during the time intervals between episodic disturbances to facilitate plant establishment, growth and survival.

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