Julie C. Pett-Ridge

Nitrogen enrichment regulates calcium sources in forests

Nitrogen (N) is a key nutrient that shapes cycles of other essential elements in forests, including calcium (Ca). When N availability exceeds ecosystem demands, excess N can stimulate Ca leaching and deplete Ca from soils. Over the long term, these processes may alter the proportion of available Ca that is derived from atmospheric deposition vs. bedrock weathering, which has fundamental consequences for ecosystem properties and nutrient supply. We evaluated how landscape variation in soil N, reflecting long-term legacies of biological N fixation, influenced plant and soil Ca availability and ecosystem Ca sources across 22 temperate forests in Oregon. We also examined interactions between soil N and bedrock Ca using soil N gradients on contrasting basaltic vs. sedimentary bedrock that differed 17-fold in underlying Ca content. We found that low-N forests on Ca-rich basaltic bedrock relied strongly on Ca from weathering, but that soil N enrichment depleted readily weatherable mineral Ca and shifted forest reliance toward atmospheric Ca. Forests on Ca-poor sedimentary bedrock relied more consistently on atmospheric Ca across all levels of soil N enrichment. The broad importance of atmospheric Ca was unexpected given active regional uplift and erosion that are thought to rejuvenate weathering supply of soil minerals. Despite different Ca sources to forests on basaltic vs. sedimentary bedrock, we observed consistent declines in plant and soil Ca availability with increasing N, regardless of the Ca content of underlying bedrock. Thus, traditional measures of Ca availability in foliage and soil exchangeable pools may poorly reflect long-term Ca sources that sustain soil fertility. We conclude that long-term soil N enrichment can deplete available Ca and cause forests to rely increasingly on Ca from atmospheric deposition, which may limit ecosystem Ca supply in an increasingly N-rich world.

Response of the nitrogen‐fixing lichen Lobaria pulmonaria to phosphorus, molybdenum, and vanadium

Nitrogen-fixing lichens (cyanolichens) are an important source of nitrogen (N) in Pacific Northwest forests, but limitation of lichen growth by elements essential for N fixation is poorly understood. To investigate how nutrient limitation may affect cyanolichen growth rates, we fertilized a tripartite cyanobacterial lichen (Lobaria pulmonaria) and a green algal non-nitrogen fixing lichen (Usnea longissima) with the micronutrients molybdenum (Mo) and vanadium (V), both known cofactors for enzymes involved in N fixation, and the macronutrient phosphorus (P). We then grew treated lichens in the field for one year in western Oregon, USA. Lichen growth was very rapid for both species and did not differ across treatments, despite a previous demonstration of P-limitation in L. pulmonaria at a nearby location. To reconcile these disparate findings, we analyzed P, Mo, and V concentrations, natural abundance δ15N isotopes, %N and change in thallus N in Lobaria pulmonaria from both growth experiments. Nitrogen levels in deposition and in lichens could not explain the large difference in growth or P limitation observed between the two studies. Instead, we provide evidence that local differences in P availability may have caused site-specific responses of Lobaria to P fertilization. In the previous experiment, Lobaria had low background levels of P, and treatment with P more than doubled growth. In contrast, Lobaria from the current experiment had much higher background P concentrations, similar to P-treated lichens in the previous experiment, consistent with the idea that ambient variation in P availability influences the degree of P limitation in cyanolichens. We conclude that insufficient P, Mo, and V did not limit the growth of either cyanolichens or chlorolichens at the site of the current experiment. Our findings point to the need to understand landscape-scale variation in P availability to cyanolichens, and its effect on spatial patterns of cyanolichen nutrient limitation and N fixation.

Nutrient feedbacks to soil heterotrophic nitrogen fixation in forests

Multiple nutrient cycles regulate biological nitrogen (N) fixation in forests, yet long-term feedbacks between N-fixation and coupled element cycles remain largely unexplored. We examined soil nutrients and heterotrophic N-fixation across a gradient of 24 temperate conifer forests shaped by legacies of symbiotic N-fixing trees. We observed positive relationships among mineral soil pools of N, carbon (C), organic molybdenum (Mo), and organic phosphorus (P) across sites, evidence that legacies of symbiotic N-fixing trees can increase the abundance of multiple elements important to heterotrophic N-fixation. Soil N accumulation lowered rates of heterotrophic N-fixation in organic horizons due to both N inhibition of nitrogenase enzymes and declines in soil organic matter quality. Experimental fertilization of organic horizon soil revealed widespread Mo limitation of heterotrophic N-fixation, especially at sites where soil Mo was scarce relative to C. Fertilization also revealed widespread absence of P limitation, consistent with high soil P:Mo ratios. Responses of heterotrophic N-fixation to added Mo (positive) and N (negative) were correlated across sites, evidence that multiple nutrient controls of heterotrophic N-fixation were more common than single-nutrient effects. We propose a conceptual model where symbiotic N-fixation promotes coupled N, C, P, and Mo accumulation in soil, leading to positive feedback that relaxes nutrient limitation of overall N-fixation, though heterotrophic N-fixation is primarily suppressed by strong negative feedback from long-term soil N accumulation.