Causes and consequences of prolonged dormancy: Why stay belowground?

Prolonged dormancy is a stage in which mature plants fail to resprout during the growing season and instead remain alive belowground. Though it is relatively common, the causes and consequences of this intriguing stage have remained elusive. In this dissertation, I investigate the causes and consequences of prolonged dormancy in a long lived perennial herb, Astragalus scaphoides. First, I use a combination of demography and ecophysiology to study the proximate mechanisms associated with prolonged dormancy. Analysis of a long-term demographic dataset indicates that both endogenous factors (e.g. age, condition, and history) and exogenous factors (e.g. climate and spatial variation) are associated with dormancy. I then investigate the association between stored resources and dormancy. My results indicate that individual plants with low levels of stored available carbon are more likely to enter prolonged dormancy. Surprisingly, individuals increased their mobile carbon concentrations while dormant, presumably by remobilizing structural carbon into mobile forms. Since stored resources integrate past conditions and performance with current state, these results can explain why some individuals remain belowground while others emerge to grow and reproduce. I used matrix models to examine the ultimate causes and consequences of prolonged dormancy. I found evidence that prolonged dormancy acts as a conservative strategy that allows plants to avoid the risk of a variable environment. Further, my results demonstrate that intermediate levels of dormancy result in the highest fitness advantage. Finally, I measured the trade-offs associated with emerging during times of environmental stress. Although plants showed remarkable physiological tolerance to stress, stress led to demographic costs. Therefore, prolonged dormancy is shown to be a beneficial strategy in a variable environment. Together, my research identifies both the proximate causes of prolonged dormancy, as well as the ultimate consequences of remaining belowground during the growing season. Therefore, my research not only identifies why some plants go dormant while others emerge, but also explains the prevalence of this intriguing life stage in the life histories of so many perennial plants

Frequent Fire Alters Nitrogen Transformations in Ponderosa Pine Stands of the Inland Northwest

Recurrent, low-severity fire in ponderosa pine (Pinus ponderosa)linterior Douglas-fir (Pseudotsuga menziesii var. glauca) forests is thought to have directly influenced nitrogen (N) cycling and availability. However, no studies to date have investigated the influence of natural fire intervals on soil processes in undisturbed forests, thereby limiting our ability to understand ecological processes and successional dynamics in this important ecosystem of the Rocky Mountain West. Here, we tested the standing hypothesis that recurrent fire in ponderosa pine/Douglas-fir forests of the Inland Northwest decreases total soil N, but increases N turnover and nutrient availability. We compared soils in stands unburned over the past 69-130 years vs. stands exposed to two or more fires over the last 130 years at seven distinct locations in two wilderness areas. Mineral soil samples were collected from each of the seven sites in June and July of 2003 and analyzed for pH, total C and N, potentially mineralizable N (PMN), and extractable NH4+, NO3, PO4-3, Ca+2, Me, and K+. Nitrogen transformations were assessed at five sites by installing ionic resin capsules in the mineral soil in August of 2003 and by conducting laboratory assays of nitrification potential and net nitrification in aerobic incubations. Total N and PMN decreased in stands subjected to multiple fires. This loss of total N and labile N was not reflected in concentrations of extractable NH4+ and NO3-. Rather, multiple fires caused an increase in NO3- sorbed on ionic resins, nitrification potential, and net nitrification in spite of the burned stands not having been exposed to fire for at least 12-17 years. Charcoal collected from a recent fire site and added to unburned soils increased nitrification potential, suggesting that the decrease of charcoal in the absence of tire may play an important role in N transformations in fire-dependent ecosystems in the long term. Interestingly, we found no consistent effect of fire frequency on extractable P or alkaline metal concentrations. Our results corroborate the largely untested hypothesis that frequent fire in ponderosa pine forests increases inorganic N availability in the long term and emphasize the need to study natural, unmanaged sites in far greater detail

Disappearing Plants: Why They Hide and How They Return

Prolonged dormancy is a life-history stage in which mature plants fail to resprout for one or more growing seasons and instead remain alive belowground. Prolonged dormancy is relatively common, but the proximate causes and consequences of this intriguing strategy have remained elusive. In this study we tested whether stored resources are associated with remaining belowground, and investigated the resource costs of remaining belowground during the growing season. We measured stored resources at the beginning and end of the growing season in Astragalus scaphoides, an herbaceous perennial in southwest Montana, USA. At the beginning of the growing season, dormant plants had lower concentrations of stored mobile carbon (nonstructural carbohydrates, NSC) than did emergent plants. Surprisingly, during the growing season, dormant plants gained as much NSC as photosynthetically active plants, an increase most likely due to remobilization of structural carbon. Thus, low levels of stored NSC were associated with remaining belowground, and remobilization of structural carbon may allow for dormant plants to emerge in later seasons. The dynamics of NSC in relation to dormancy highlights the ability of plants to change their own resource status somewhat independently of resource assimilation, as well as the importance of considering stored resources in understanding plant responses to the environment

Arl3 regulates a transport system for farnesylated cargo

Arl3 is a small G-protein that is found exclusively in ciliated organisms. In addition, knocking out of Arl3 results in a plethora of ciliopathies. Arl3 is known to bind the photoreceptor (specialized cilia) specific PDE delta subunit (PDE6D), which in turn bind to prenylated proteins. The significance of this interaction and the function of Arl3 in cilia are poorly understood. Here in this study, by solving the crystal structure of a fully modified prenylated (farnesylated) Rheb in complex with PDE6D and comparing it to a structure of PDE6D in complex with the Arl3 homologue Arl2, we show that Arl3 is an allosteric regulator of PDE6D. Arl3, in a nucleotide dependent manner, releases the farnesylated cargo bound to PDE6D. We explain the molecular mechanism of this release and we further verify the mechanism in vitro and by live cell imaging. Based on this study we hypothesize that Arl3 regulate the targeting of prenylated cargo in and out the cilia

Optimization of the All-D peptide D3 for Aβ oligomer elimination

The aggregation of amyloid-{beta} (A{beta}) is postulated to be the crucial event in Alzheimer's disease (AD). In particular, small neurotoxic A{beta} oligomers are considered to be responsible for the development and progression of AD. Therefore, elimination of thesis oligomers represents a potential causal therapy of AD. Starting from the well-characterized d-enantiomeric peptide D3, we identified D3 derivatives that bind monomeric A{beta}. The underlying hypothesis is that ligands bind monomeric A{beta} and stabilize these species within the various equilibria with A{beta} assemblies, leading ultimately to the elimination of A{beta} oligomers. One of the hereby identified d-peptides, DB3, and a head-to-tail tandem of DB3, DB3DB3, were studied in detail. Both peptides were found to: (i) inhibit the formation of Thioflavin T-positive fibrils; (ii) bind to A{beta} monomers with micromolar affinities; (iii) eliminate A{beta} oligomers; (iv) reduce A{beta}-induced cytotoxicity; and (v) disassemble preformed A{beta} aggregates. The beneficial effects of DB3 were improved by DB3DB3, which showed highly enhanced efficacy. Our approach yielded A{beta} monomer-stabilizing ligands that can be investigated as a suitable therapeutic strategy against AD

Testing the Trait-Based Community Framework: Do Functional Traits Predict Competitive Outcomes?

Plant traits can be used to understand a range of ecological processes, including competition from invasive species. The extent to which native and invasive species are competing via limiting similarity or trait hierarchies has important implications for the management of invaded communities. We screened 47 native species that co-occur with Festuca perennis, a dominant invader in California serpentine grassland, for traits pertaining to resource use and acquisition. We then grew F. perennis with ten species spanning a range of functional similarity in pairwise competition trials. Functionally similar species did not have a strong adverse effect on F. perennis performance as would be expected by limiting similarity theory. Phylogenetic relatedness, which may integrate a number of functional traits, was also a poor predictor of competitive outcome. Instead, species with high specific root length, low root to shoot biomass ratio, and low leaf nitrogen concentration were more effective at suppressing the growth of F. perennis. Our results suggest that fitness differences (i.e., trait hierarchies) may be more important than niche differences (i.e., limiting similarity) in structuring competitive outcomes in this system and may be a promising approach for the restoration of invaded systems

Regulatory coiled-coil domains promote head-to-head assemblies of AAA+ chaperones essential for tunable activity control

Ring-forming AAA+ chaperones exert ATP-fueled substrate unfolding by threading through a central pore. This activity is potentially harmful requiring mechanisms for tight repression and substrate-specific activation. The AAA+ chaperone ClpC with the peptidase ClpP forms a bacterial protease essential to virulence and stress resistance. The adaptor MecA activates ClpC by targeting substrates and stimulating ClpC ATPase activity. We show how ClpC is repressed in its ground state by determining ClpC cryo-EM structures with and without MecA. ClpC forms large two-helical assemblies that associate via head-to-head contacts between coiled-coil middle domains (MDs). MecA converts this resting state to an active planar ring structure by binding to MD interaction sites. Loss of ClpC repression in MD mutants causes constitutive activation and severe cellular toxicity. These findings unravel an unexpected regulatory concept executed by coiled-coil MDs to tightly control AAA+ chaperone activity