Rapid biodiversity declines in both ungrazed and intensely grazed exotic grasslands

Exotic-dominated ecosystems with low diversity are becoming increasingly common. It remains unclear, though, whether differences between native and exotic species (driver model), or changes in disturbances or resources (passenger model), allow exotics to become competitive dominants. In our field experiment, plant species origin (native or exotic), cattle grazing (ungrazed or intensely grazed once), and species composition treatments were fully crossed and randomly assigned to four-species mixtures and monocultures of grassland plants. We found that biodiversity declined more rapidly in exotic than in native species mixtures, regardless of our grazing disturbance treatment. Early declines in species evenness (i.e., increases in dominance) led to subsequent declines in species richness (i.e., local extinctions) in exotic mixtures. Specifically, Simpson’s diversity was 29% lower after 1 year, and species richness was 15% lower after 3 years, in exotic than in native mixtures. These rapid biodiversity declines in exotic mixtures were partly explained by decreased complementarity (i.e., niche partitioning and facilitation), presumably because exotic species lack the coevolutionary history that can lead to complementarity and coexistence in native communities. Thus, our results suggest that exotic species can drive biodiversity declines in the presence or absence of a grazing disturbance, partly because exotic species interactions differ from native species interactions. This implies that restoring plant biodiversity in grasslands may require removal of exotic species, in addition to disturbance management.     

Invasive plants negatively impact native,but not exotic,animals

Despite our growing understanding of the impacts of invasive plants on ecosystem structure and function, important gaps remain, including whether native and exotic species respond differently to plant invasion. This would elucidate basic ecological interactions and inform management. We performed a meta‐analytic review of the effects of invasive plants on native and exotic resident animals. We found that invasive plants reduced the abundance of native, but not exotic, animals. This varied by animal phyla, with invasive plants reducing the abundance of native annelids and chordates, but not mollusks or arthropods. We found dissimilar impacts among “wet” and “dry” ecosystems, but not among animal trophic levels. Additionally, the impact of invasive plants increased over time, but this did not vary with animal nativity. Our review found that no studies considered resident nativity differences, and most did not identify animals to species. We call for more rigorous studies of invaded community impacts across taxa, and most importantly, explicit consideration of resident biogeographic origin. We provide an important first insight into how native and exotic species respond differently to invasion, the consequences of which may facilitate cascading trophic disruptions further exacerbating global change consequences to ecosystem structure and function.     

Changes in vegetation states in grazed and ungrazed Mackenzie Basin grasslands, New Zealand, 1990-2000

Changes in vegetation from 1990 to 2000 were examined at 10 high country localities, representing four grassland types: fescue tussock (Festuca novae-zelandiae), snow tussock (Chionochloa rigida), red tussock (C. rubra), and silver tussock (Poa cita). At each locality, three treatments were established: ambient sheep+rabbit grazing, rabbit grazing only, and no grazing. The mutivariate methods of classification and ordination were used on individual-quadrat cover data to define vegetation states and to examine transitions between them over time. Vegetation states in quadrats already dominated by Hieracium pilosella(> 50% cover) in 1990 showed little change in species composition regardless of grassland type and grazing treatment. In fescue tussock grassland, H. pilosellaincreased regardless of grazing treatment in states with low initial H. pilosellacover (< 5%), while the cover of Carex colensoi, Aira caryophyllea and Rumex acetosella decreased. In the single silver tussock locality, Poa citadecreased markedly in the ungrazed treatment as adventive species such as Dactylis glomerataand Echium vulgare increased. However, Poa citaalso decreased, probably due to drought, in the grazed treatment. Snow tussock and red tussock grassland states were more stable than those in short tussock grasslands, but there was also a general trend towards increasing H. pilosellacover in intertussock vegetation regardless of treatment. However, at one snow tussock locality, transitions from H. piloselladominated to C. rigida-dominated states occurred in ungrazed quadrats, while the reverse occurred in grazed vegetation. Implications for the management of tussock grasslands for conservation are discussed.     

Increasing precipitation event size increases aboveground net primary productivity in a semi-arid grassland

Water availability is the primary constraint to aboveground net primary productivity (ANPP) in many terrestrial biomes, and it is an ecosystem driver that will be strongly altered by future climate change. Global circulation models predict a shift in precipitation patterns to growing season rainfall events that are larger in size but fewer in number. This “repackaging” of rainfall into large events with long intervening dry intervals could be particularly important in semi-arid grasslands because it is in marked contrast to the frequent but small events that have historically defined this ecosystem. We investigated the effect of more extreme rainfall patterns on ANPP via the use of rainout shelters and paired this experimental manipulation with an investigation of long-term data for ANPP and precipitation. Experimental plots (n = 15) received the long-term (30-year) mean growing season precipitation quantity; however, this amount was distributed as 12, six, or four events applied manually according to seasonal patterns for May–September. The long-term mean (1940–2005) number of rain events in this shortgrass steppe was 14 events, with a minimum of nine events in years of average precipitation. Thus, our experimental treatments pushed this system beyond its recent historical range of variability. Plots receiving fewer, but larger rain events had the highest rates of ANPP (184 ± 38 g m⁻²), compared to plots receiving more frequent rainfall (105 ± 24 g m⁻²). ANPP in all experimental plots was greater than long-term mean ANPP for this system (97 g m⁻²), which may be explained in part by the more even distribution of applied rain events. Soil moisture data indicated that larger events led to greater soil water content and likely permitted moisture penetration to deeper in the soil profile. These results indicate that semi-arid grasslands are capable of responding immediately and substantially to forecast shifts to more extreme precipitation patterns.     

Temporal coherence of aboveground net primary productivity in mesic grasslands

Synchrony in ecological variables over wide geographic areas suggests that large‐scale environmental factors drive the structure and function of ecosystems and override more local‐scale environmental variation. Described also as coherence, this phenomenon has been documented broadly in the ecological literature and has recently received increasing attention as scientists attempt to quantify the impacts of global changes on organisms and their habitats. Using a mesic grassland site in North America, we assessed coherence in ecosystem function by quantifying similarity in aboveground net primary production (ANPP) dynamics in 48 permanent sampling locations (PSLs) over a 16‐yr period. Our primary objective was to characterize coherence across a broad geographic region (with similar ecosystem structure and function), and we hypothesized that precipitation and a similar fire frequency would strengthen coherence between PSLs. All 48 PSLs at our site (Konza Prairie Biological Station, Manhattan, KS, USA; KPBS) were exposed to a similar regional driver of ANPP (precipitation); however, local drivers (including differences in fire frequency and soil depth at different topographic positions) varied strongly among individual PSLs. For the purpose of this assessment, the watershed‐level experimental design of KPBS was considered a model, which represented different fire management strategies across the Great Plains Region. Our analyses revealed a site‐level (KPBS) coherence in ANPP dynamics of 0.53 for the period of 1984–1999. Annual fire enhanced coherence among PSLs to 0.76, whereas less frequent fire (fire exclusion or a 4‐yr fire return interval) failed to further increase coherence beyond that of the KPBS site level. Soil depth also strongly influenced coherence among PSLs with shallow soils at upland sites showing strong coherence across fire regimes and annually burned uplands closely linked to annual precipitation dynamics. The lack of coherence in ecosystem function in PSLs with deep soils and low fire frequencies suggests that conservation and management efforts will need to be more location specific in such areas where biotic interactions may be more important than regional abiotic drivers.     

An invasive plant provides refuge to native plant species in an intensely grazed ecosystem

Invasion by exotic plant species and herbivory can individually alter native plant species diversity, but their interactive effects in structuring native plant communities remain little studied. Many exotic plant species escape from their co-evolved specialized herbivores in their native range (in accordance with the enemy release hypothesis). When these invasive plants are relatively unpalatable, they may act as nurse plants by reducing herbivore damage on co-occurring native plants, thereby structuring native plant communities. However, the potential for unpalatable invasive plants to structure native plant communities has been little investigated. Here, we tested whether presence of an unpalatable exotic invader Opuntia ficus-indica was associated with the structure of native plant communities in an ecosystem with a long history of grazing by ungulate herbivores. Along 17 transects (each 1000 m long), we conducted a native vegetation survey in paired invaded and uninvaded plots. Plots that harboured O. ficus-indica had higher native plant species richness and Shannon–Wiener diversity H′ than uninvaded plots. However, mean species evenness J was similar between invaded and uninvaded plots. There was no significant correlation between native plant diversity and percentage plot cover by O. ficus-indica. Presence of O. ficus-indica was associated with a compositional change in native community assemblages between paired invaded and uninvaded plots. Although these results are only correlative, they suggest that unpalatable exotic plants may play an important ecological role as refugia for maintenance of native plant diversity in intensely grazed ecosystems.     

Non-random co-occurrence of native and exotic plant species in Mediterranean grasslands

Invasion by exotic species in Mediterranean grasslands has determined assembly patterns of native and introduced species, knowledge of which provides information on the ecological processes underlying these novel communities. We considered grasslands from Spain and Chile. For each country we considered the whole grassland community and we split species into two subsets: in Chile, species were classified as natives or colonizers (i.e. exotics); in Spain, species were classified as exclusives (present in Spain but not in Chile) or colonizers (Spanish natives and exotics into Chile). We used null models and co-occurrence indices calculated in each country for each one of 15 sites distributed along a precipitation gradient and subjected to similar silvopastoral exploitation. We compared values of species co-occurrence between countries and between species subsets (natives/colonizers in Chile; exclusives/colonizers in Spain) within each country and we characterised them according to climatic variables. We hypothesized that: a) the different coexistence time of the species in both regions should give rise to communities presenting a spatial pattern further from random in Spain than in Chile, b) the co-occurrence patterns in the grasslands are affected by mesoclimatic factors in both regions. The patterns of co-occurrence are similar in Spain and Chile, mostly showing a spatial pattern more segregated than expected by random. The colonizer species are more segregated in Spain than in Chile, possibly determined by the longer residence time of the species in the source area than in the invaded one. The segregation of species in Chile is related to water availability, being species less segregated in habitat with greater water deficit; in Spain no relationship with climatic variables was found. After an invasion process, our results suggest that the possible process of alteration of the original Chilean communities has not prevented the assembly between the native and colonizer species together.     

Breakdown of the species-area relationship in exotic but not in native forest patches

We studied the pattern of bird species richness in native and exotic forest patches in Hungary. We hypothesized that species-area relationship will depend on forest naturalness, and on the habitat specialization of bird species. Therefore, we expected strong species-area relationship in native forest patches and forest bird species, and weaker relationship in exotic forest patches containing generalist species. We censused breeding passerine bird communities three times in 13 forest patches with only native tree species, and 14 with only exotic trees in Eastern Hungary in 2003. Although most bird species (92%) of the total of 41 species occurred in both exotic and native forests, the species-area relationship was significant for forest specialist, but not for generalist species in the native forests. No relationship between bird species and area was found for either species group in the forest with exotic tree species. The comparison of native versus exotic forest patches of similar sizes revealed that only large (>100 ha) native forests harbor higher bird species richness than exotic forests for the forest specialist bird species. There is no difference between small and medium forest patches and in richness of generalist species. Thus, the species-area relationship may diminish in archipelago of exotic habitat patches and/or for habitat generalist species; this result supports the warning that the extension of exotic habitats have been significantly contributing to the decline of natural community patterns.     

Spread of exotic grass in grazed native grass pastures and responses of insect communities

Exotic grasses are widely established across the Southeastern United States for livestock forage, resulting in the structural and compositional simplification of grasslands. Replacing exotic forages with native warm‐season grasses (NWSG) could benefit insects due to increased complexity of plant structure and composition, but livestock grazing also may facilitate spread of remnant exotic grasses such as bermudagrass (Cynodon dactylon) by reducing height and coverage of NWSG. We investigated these relationships among 12 operational‐scale pastures (6.4–10.5 ha) in Mississippi, U.S.A., during May–July (2011–2012). We quantified changes in bermudagrass coverage from one treatment of grazed exotic forages and three treatments of recently established NWSG, including a grazed mixed NWSG polyculture, a grazed Indian grass (Sorghastrum nutans) monoculture to evaluate the effects of stand‐type richness among NWSG pastures, and a non‐grazed NWSG polyculture to evaluate the effects of grazing. We also assessed responses of two insect orders, Orthoptera and Hemiptera, to treatment and bermudagrass coverage. We estimated a 101–190% average increase in coverage of bermudagrass in grazed native grass pastures (NWSG polyculture and Indian grass monoculture), but not in non‐grazed NWSG, suggesting that grazing facilitated the spread of this grass. Composition of Orthopteran and Hemipteran communities was correlated with bermudagrass coverage, and inter‐year differences in composition for both communities in grazed mixed NWSG, and for Hemiptera in grazed Indian grass, corresponded with increasing bermudagrass coverage in those treatments. Our results suggest that incomplete eradication of exotic forages prior to establishment of NWSG may be exacerbated by grazing, which could then impact stand condition and insect communities.     

Changes in light- and nitrogen-use and in aboveground biomass allocation patterns along productivity gradients in grasslands

Light- and nitrogen-use change was examined along productivity gradients in natural grasslands at Laelatu, western Estonia, both at community level and in most abundant species. Aboveground biomass (M) ranged from 341 to 503 g m^?2 in wet (W) and from 248 to 682 g m^?2 in dry (D) community. Aboveground leaf area ratio (aLAR) decreased with rising M in D site, while it increased in W site. In a high-aLAR W community (significantly higher compared to D), adjustment of leaf morphology through an increase in specific leaf area is responsible for an increase in aLAR with rising productivity. In low-aLAR stand, by contrast, adjustment of biomass allocation due to decrease in aboveground leaf mass fraction is primarily responsible for the tendency of aLAR to decline. In conclusion, a decrease in aLAR is not a universal response to increasing M. We hypothesise that there exists an optimum of light acquisition efficiency (Φ_M) along a productivity gradient independent of community type. Aboveground nitrogen-use efficiency (aNUE) decreased in high-aLAR, W community with increasing M, while in low-aLAR, D site, there was no relationship along a gradient, although aNUE increased along six plots dominated by graminoids. A trade-off was established between leaf nitrogen content per unit leaf area (N _A) and aLAR.     

Below‐ground biomass and productivity of a grazed site and a neighbouring ungrazed exclosure in a grassland in central Argentina

Abstract We estimated the below‐ground net plant productivity (BNPP) of different biomass components in an intensively and continuously 45‐ha grazed site and in a neighbouring exclosure ungrazed for 16 years for a natural mountain grassland in central Argentina. We measured approximately twice as much dead below‐ground biomass in the grazed site as in the ungrazed site, with a strong concentration of total below‐ground biomass towards the upper 10 cm of the soil layer in both sites. The main contribution to total live biomass was accounted for by very fine (<0.5 mm) and fine roots (0.5–1.0 mm) both at the grazed (79%) and at the ungrazed (81%) sites. We measured more dead biomass for almost all root components, more live biomass of rhizomes, tap roots and bulbs, and less live biomass of thicker roots (>1 mm) in the grazed site. The seasonal variation of total live below‐ground biomass mainly reflected climate, with the growing season being limited to the warmer and wetter portion of the year, but such variation was higher in the grazed site. Using different methods of estimation of BNPP, we estimated maximum values of 1241 and 723 g m^?2 year^?1 for the grazed and ungrazed sites, respectively. We estimated that very fine root productivity was almost twice as high at the grazed site as at the ungrazed one, despite the fact that both sites had similar total live biomass, and root turnover rate was twofold at the grazed site.     

Local soil,but not commercial AMF inoculum,increases native and non‐native grass growth at a mine restoration site

Soil communities are often degraded in mined sites, and facilitating the recovery of soil mutualists such as arbuscular mycorrhizal fungi (AMF) may assist with the restoration of native plants. At a grassland mine restoration site, I compared a commercial AMF inoculum with soil collected from beneath native grasses as a source of inoculum, as well as a control treatment. Field plots were broadcast‐inoculated and seeded with native grasses, and biomass of native and non‐native species was measured in three consecutive years. In addition, greenhouse‐grown seedlings of a native bunchgrass (Stipa pulchra) were inoculated with similar treatments, transplanted into the field, and assessed after 18 months. When broadcast inoculation was used, the local soil inoculum tended to increase non‐native grass biomass, and marginally decreased non‐native forb biomass in the second year of study, but did not significantly affect native grass biomass. Broadcast commercial inoculum had no detectable effects on biomass of any plant group. Stipa pulchra transplants had greater N content and mycorrhizal colonization, and marginally higher shoot mass and K content, when pre‐inoculated with local soil (relative to controls). Pre‐inoculation with commercial AMF increased AMF colonization of the S. pulchra transplants, but did not significantly affect biomass or nutrient content. The findings indicate that at this site, the use of local soil as an inoculum had greater effects on native and non‐native plants than the commercial product used. In order to substantially increase native grass performance, inoculation of transplanted plugs may be one potential strategy.     

Local adaptation of mycorrhizae communities changes plant community composition and increases aboveground productivity

Soil microbial communities can have an important role in the adaptation of plants to their local abiotic soil conditions and in mediating plant responses to environmental stress. This has been clearly demonstrated for individual plant species, but it is unknown how locally adapted microbes may affect plant communities. It is possible that the adaptation of microbial communities to local conditions can shape plant community composition. Additionally, it is possible that the effects of locally adapted microorganisms on individual plant species could be altered by co-occurring plant species. We tested these possibilities in plant community mesocosms with soils and mycorrhizal fungi (AMF) from three locations. We found that plant community biomass responded positively to local adaptation of AMF to soil conditions. Plant community composition also changed in response to local adaptation of AMF. Unexpectedly, the strongest benefits of locally adapted AMF went to early successional plant species that have the highest relative growth rates and the lowest responsiveness to the presence of AMF. Late successional plants that responded positively overall to the presence of AMF were often suppressed in communities with local AMF, perhaps because of strong competition from fast growing plant species. These results show that local adaptation of soil microbial communities can shape plant community composition, and the benefits that plants derive from locally adapted microorganisms can be reshaped by the competitive context in which these associations occur.     

Soil ecosystem function under native and exotic plant assemblages as alternative states of successional grasslands

Old fields often become dominated by exotic plants establishing persistent community states. Ecosystem functioning may differ widely between such novel communities and the native-dominated counterparts. We evaluated soil ecosystem attributes in native and exotic (synthetic) grass assemblages established on a newly abandoned field, and in remnants of native grassland in the Inland Pampa, Argentina. We asked whether exotic species alter soil functioning through the quality of the litter they shed or by changing the decomposition environment. Litter decomposition of the exotic dominant Festuca arundinacea in exotic assemblages was faster than that of the native dominant Paspalum quadrifarium in native assemblages and remnant grasslands. Decomposition of a standard litter (Triticum aestivum) was also faster in exotic assemblages than in native assemblages and remnant grasslands. In a common garden, F. arundinacea showed higher decay rates than P. quadrifarium, which reflected the higher N content and lower C:N of the exotic grass litter. Soil respiration rates were higher in the exotic than in the native assemblages and remnant grasslands. Yet there were no significant differences in soil N availability or net N mineralization between exotic and native assemblages. Our results suggest that exotic grass dominance affected ecosystem function by producing a more decomposable leaf litter and by increasing soil decomposer activity. These changes might contribute to the extended dominance of fast-growing exotic grasses during old-field succession. Further, increased organic matter turnover under novel, exotic communities could reduce the carbon storage capacity of the system in the long term.     

Soil arthropod composition differs between old-fields dominated by exotic plant species and remnant native grasslands

Secondary succession after agriculture abandonment (old-fields) is mostly dominated by exotic grass species. Non-native plant invasions may alter soil fauna, potentially inducing plant-soil feedbacks. Despite their importance in nutrient cycling and plant-soil interactions, meso and macrofauna received less attention than bacteria or fungi. Here we compared the composition of the soil arthropod community in native remnants and plant exotic-dominated old-fields grasslands in the Inland Pampa, Argentina. We sampled independent remnants and old-field grassland plots within a 100 km² agricultural landscape to test the hypothesis that the abundance of soil arthropod organisms is related to the quality of the plant biomass, whereas the diversity of the soil biota is related to plant species richness, resulting in a different soil biota composition because of differing plant communities. When compared to non-invaded remnant grasslands, soil activity and soil food-web characteristics of the old-fields sites included: 1. Higher total arthropod abundance, particularly of Isopoda, Pseudoescorpionida and Blattaria; 2. Lower abundance of Hymenoptera and Enthomobryomorpha (Collembola); 3. Lower diversity, and evenness, but similar richness of soil organisms orders; 4. Higher soil respiration rates and soil temperature; and 5. Higher total soil N and K⁺content, but lower soil P content. These results illustrate that soil arthropod composition can vary widely within grasslands patches depending on plant species composition. Also, the more diverse plant community of remnant grasslands supports a more diverse soil biota, although soil activity is slower. Our results support the strong linkage between plant community and soil arthropod composition and suggest that changes in soil biota composition might promote plant-soil feedback interactions inducing the persistence of these alternative grassland states in new agricultural human-modified landscapes.     

Predicting impact of freshwater exotic species on native biodiversity: Challenges in spatial scaling

Abstract Global homogenization of biota is underway through worldwide introduction and establishment of nonindigenous (exotic) species. Freshwater ecologists should devote more attention to exotic species for two reasons. First, exotics provide an opportunity to test hypotheses about what characteristics of species or habitats are related to successful establishment or invasibility, respectively. Second, predicting which species will cause large ecological change is an important challenge for natural resource managers. Rigorous statistical relationships linking species characteristics to probability of establishment or of causing ecological impacts are needed. In addition, it is important to know how reliable different sorts of experiments are in guiding predictions. We address this issue with different spatial scales of experiments testing the impact of two predators on native snail assemblages in northern Wisconsin USA lakes: an exotic crayfish, the rusty crayfish (Orconectes rusticus); and a native fish predator, the pumpkinseed sunfish (Lepomis gibossus). For the crayfish, laboratory experiments, a field cage experiment, and a snapshot survey of 21 lakes gave consistent results: the crayfish reduced abundance and species richness of native snails. Laboratory and field experiments suggested that pumpkinseed sunfish should have a similar impact, but the lake survey suggested little impact. Unfortunately, no algorithms exist to guide scaling up from small-scale experiments to the whole-lake, long-term management scale. To protect native biodiversity, management of freshwater exotic species should be targeted on lakes or drainages that are both vulnerable to colonization by an exotic, and that harbour endemic species. Management should focus on preventing introduction because eradication after establishment is usually not possible.     

Grouping plant species by shared native range, and not by native status, predicts response to an exotic herbivore

Differences among exotic species can be as large as differences between native and exotic species. Typically, however, only the distinction between native and exotic is made when predicting responses in a community. In this paper, I examine the response of plant species to experimental disturbance and exclusion of invasive European rabbits (Oryctolagus cuniculus) in a grassland community with exotic plants originating from five continents. I explore group responses based on native status, shared native range with rabbits, having a congener from the native range of rabbits, life-history (e.g., annual), and life-form (e.g., grass). Individual species responses to rabbits were idiosyncratic, but group responses were predicted by continent of origin, not native status. Native status did predict response to disturbance with almost uniform responses within groups. Exotic species, regardless of origin, were positively affected by disturbance. Native species, in contrast, were negatively affected by disturbance. These results suggest that grouping plant species by native status is valid for questions of disturbance, but when analyzing outcomes of interactions, factors other than native status, such as shared evolutionary history, should be considered.     

Increasing oxidative stress with molsidomine increases blood pressure in genetically hypertensive rats but not normotensive controls

Spontaneously hypertensive rats (SHR) have a higher level of oxidative stress and exhibit a greater depressor response to a superoxide scavenger, tempol, than normotensive Wistar-Kyoto rats (WKY). This study determined whether an increase in oxidative stress with a superoxide/NO donor, molsidomine, would amplify the blood pressure in SHR. Male SHR and WKY were given molsidomine (30 mg.kg(-1).day(-1)) or vehicle (0.01% ethanol) for 1 wk, and blood pressure, renal hemodynamics, nitrate and nitrite excretion (NOx), renal superoxide production, and expression of renal antioxidant enzymes, Mn- and Cu,Zn-SOD, catalase, and glutathione peroxidase (GPx), were measured. Renal superoxide and NOx were higher in control SHR than in WKY. Molsidomine increased superoxide by approximately 35% and NOx by 250% in both SHR and WKY. Mean arterial blood pressure (MAP) was also higher in control SHR than WKY. Molsidomine increased MAP by 14% and caused renal vasoconstriction in SHR but reduced MAP by 16%, with no effect on renal hemodynamics, in WKY. Renal expression of Mn- and Cu,Zn-SOD was not different between SHR and WKY, but expression of catalase and GPx were approximately 30% lower in kidney of SHR than WKY. The levels of Mn- and Cu,Zn-SOD were not increased with molsidomine in either WKY or SHR. Renal catalase and GPx expression was increased by 300-400% with molsidomine in WKY, but there was no effect in SHR. Increasing oxidative stress elevated blood pressure further in SHR but not WKY. WKY are likely protected because of higher bioavailable levels of NO and the ability to upregulate catalase and GPx.