Trade-offs between deer herbivory and nitrogen competition alter grassland forb composition.

Two of the major factors that control the composition of herbaceous plant communities are competition for limiting soil resources and herbivory. We present results from a 14-year full factorial experiment in a tallgrass prairie ecosystem that crossed nitrogen (N) addition with fencing to exclude white-tailed deer, Odocoileus virginianus, from half the plots. Deer presence was associated with only modest decreases in aboveground plant biomass (14% decrease; -45 ± 19 g m-2) with no interaction with N addition. N addition at 5.44 and 9.52 g N m-2 year-1 led to increases in biomass. There were weak increases in species richness associated with deer presence, but only for no or low added N (1 and 2 g N m-2 year-1). However, the presence of deer greatly impacted the abundances of some of the dominant perennial forb species, but not the dominant grasses. Deer presence increased the abundance of the forb Artemisia ludoviciana by 34 ± 12 SE g m-2 (94%) and decreased the forb Solidago rigida by 32 ± 13 SE g m-2 (79%). We suggest that these changes may have resulted from trade-offs in plant competitive ability for soil N versus resistance to deer herbivory. Field observations suggest deer acted as florivores, mainly consuming the flowers of susceptible forb species. The preferential consumption of flowers of forbs that seem to be superior N competitors appears to create an axis of interspecific niche differentiation. The overpopulation of white-tailed deer in many tallgrass reserves likely structures the abundance of forb species.

Plant chemical traits define functional and phylogenetic axes of plant biodiversity.

To determine which types of plant traits might better explain ecosystem functioning and plant evolutionary histories, we compiled 42 traits for each of 15 perennial species in a biodiversity experiment. We used every possible combination of three traits to cluster species. Across these 11,480 combinations, clusters generated using tissue %Ca, %N and %K best mapped onto phylogeny. Moreover, for the 15 best combinations of three traits, 82% of traits were chemical, 16% morphological and 2% metabolic. The diversity-dependence of ecosystem productivity was better explained by the %Ca, %N and %K clusters: compared to adding a new species at random, adding a species from an absent cluster/clade better-explained gains in productivity. Species number impacted productivity only when all clusters were present. Our results suggest that tissue elemental chemistry might be more phylogenetically conserved and more strongly related to ecosystem functioning than commonly measured morphological and physiological traits, a possibility that merits exploration.

Might field experiments also be inadvertent metacommunities?

Metacommunity theory predicts that the composition and diversity of a site depend on its characteristics and those of its neighborhood. Dispersal between plots in a field experiment could link responses observed in a focal plot to both its treatment and those of its neighbors. However, the diversity, composition, and treatments of neighboring plots are rarely included in analyses of experimental treatments. We analyzed a spatially gridded grassland nitrogen addition experiment and found that plant species richness and the composition of focal plots were influenced not just by their nitrogen treatment but also by the number of species in neighboring plots and their abundances. For each additional species in a focal plot's neighborhood, the species richness of the focal plot increased by 0.30 species. Control plots had a significant loss of species, at a rate of ~0.23 species per year during the 23-year experiment, but only when their neighborhoods had low species richness. Changes in the abundance of the three dominant species depended both on the nitrogen treatment of a focal plot and on their abundance in adjacent plots. Our analyses suggested that both the experimental nitrogen treatments and metacommunity processes codetermined plant species richness and plant species' abundances. Our findings suggested that analyzing many traditional field experiments with a metacommunity perspective may reveal a confounding of experimental treatments and provide empirical data to test metacommunity theory.

Plant biodiversity and the regeneration of soil fertility.

Fertile soils have been an essential resource for humanity for 10,000 y, but the ecological mechanisms involved in the creation and restoration of fertile soils, and especially the role of plant diversity, are poorly understood. Here we use results of a long-term, unfertilized plant biodiversity experiment to determine whether biodiversity, especially plant functional biodiversity, impacted the regeneration of fertility on a degraded sandy soil. After 23 y, plots containing 16 perennial grassland plant species had, relative to monocultures of these same species, ∼30 to 90% greater increases in soil nitrogen, potassium, calcium, magnesium, cation exchange capacity, and carbon and had ∼150 to 370% greater amounts of N, K, Ca, and Mg in plant biomass. Our results suggest that biodiversity, likely in combination with the increased plant productivity caused by higher biodiversity, led to greater soil fertility. Moreover, plots with high plant functional diversity, those containing grasses, legumes, and forbs, accumulated significantly greater N, K, Ca, and Mg in the total nutrient pool (plant biomass and soil) than did plots containing just one of these three functional groups. Plant species in these functional groups had trade-offs between their tissue N content, tissue K content, and root mass, suggesting why species from all three functional groups were essential for regenerating soil fertility. Our findings suggest that efforts to regenerate soil C stores and soil fertility may be aided by creative uses of plant diversity.

Diversity-dependent soil acidification under nitrogen enrichment constrains biomass productivity.

In most plant communities, the net effect of nitrogen enrichment is an increase in plant productivity. However, nitrogen enrichment also has been shown to decrease species richness and to acidify soils, each of which may diminish the long-term impact of nutrient enrichment on productivity. Here we use a long-term (20 year) grassland plant diversity by nitrogen enrichment experiment in Minnesota, United States (a subexperiment within the BioCON experiment) to quantify the net impacts of nitrogen enrichment on productivity, including its potential indirect effects on productivity via changes in species richness and soil pH over an experimental diversity gradient. Overall, we found that nitrogen enrichment led to an immediate positive increment in productivity, but that this effect became nonsignificant over later years of the experiment, with the difference in productivity between fertilized and unfertilized plots decreasing in proportion to nitrogen addition-dependent declines in soil pH and losses of plant diversity. The net effect of nitrogen enrichment on productivity could have been 14.5% more on average over 20 years in monocultures if not for nitrogen-induced decreases in pH and about 28.5% more on average over 20 years in 16 species communities if not for nitrogen-induced species richness losses. Together, these results suggest that the positive effects of nutrient enrichment on biomass production can diminish in their magnitude over time, especially because of soil acidification in low diversity communities and especially because of plant diversity loss in initially high diversity communities.

Creatine supplementation prevents the accumulation of fat in the livers of rats fed a high-fat diet.

The aim of the present study was to examine the effects of creatine supplementation on liver fat accumulation induced by a high-fat diet in rats. Rats were fed 1 of 3 different diets for 3 wk: a control liquid diet (C), a high-fat liquid diet (HF), or a high-fat liquid diet supplemented with creatine (HFC). The C and HF diets contained, respectively, 35 and 71% of energy derived from fat. Creatine supplementation involved the addition of 1% (wt:v) of creatine monohydrate to the liquid diet. The HF diet increased total liver fat concentration, liver TG, and liver TBARS and decreased the hepatic S-adenosylmethionine (SAM) concentration. Creatine supplementation normalized all of these perturbations. Creatine supplementation significantly decreased the renal activity of l-arginine:glycine amidinotransferase and plasma guanidinoacetate and prevented the decrease in hepatic SAM concentration in rats fed the HF diet. However, there was no change in either the phosphatidylcholine:phosphatidylethanolamine (PE) ratio or PE N-methyltransferase activity. The HF diet decreased mRNA for PPARα as well as 2 of its targets, carnitine palmitoyltransferase and long-chain acylCoA dehydrogenase. Creatine supplementation normalized these mRNA levels. In conclusion, creatine supplementation prevented the fatty liver induced by feeding rats a HF diet, probably by normalization of the expression of key genes of ß-oxidation.