MRI-derived Articular Cartilage Strains Predict Patient-Reported Outcomes Six Months Post Anterior Cruciate Ligament Reconstruction.

Key terms: Multicontrast and Multiparametric, Magnetic Resonance Imaging, Osteoarthritis, Functional Biomechanical Imaging, Knee Joint Degeneration What is known about the subject: dualMRI has been used to quantify strains in a healthy human population in vivo and in cartilage explant models. Previously, OA severity, as determined by histology, has been positively correlated to increased shear and transverse strains in cartilage explants. What this study adds to existing knowledge: This is the first in vivo use of dualMRI in a participant demographic post-ACL reconstruction and at risk for developing osteoarthritis. This study shows that dualMRI-derived strains are more significantly correlated with patient-reported outcomes than any MRI relaxometry metric. Background: Anterior cruciate ligament (ACL) injuries lead to an increased risk of osteoarthritis, characterized by altered cartilage tissue structure and function. Displacements under applied loading by magnetic resonance imaging (dualMRI) is a novel MRI technique that can be used to quantify mechanical strain in cartilage while undergoing a physiological load. Purpose: To determine if strains derived by dualMRI and relaxometry measures correlate with patient-reported outcomes at six months post unilateral ACL reconstruction. Study Design: Cohort study. Methods: Quantitative MRI (T2, T2*, T1ρ) measurements and transverse, axial, and shear strains were quantified in the medial articular tibiofemoral cartilage of 35 participants at six-months post unilateral ACL reconstruction. The relationships between patient-reported outcomes (WOMAC, KOOS, MARS) and all qMRI relaxation times were quantified using general linear mixed-effects models. A combined best-fit multicontrast MRI model was then developed using backwards regression to determine the patient features and MRI metrics that are most predictive of patient-reported outcome scores. Results: Higher femoral strains were significantly correlated with worse patient-reported functional outcomes. Femoral shear and transverse strains were positively correlated with six-month KOOS and WOMAC scores, after controlling for covariates. No relaxometry measures were correlated with patient-reported outcome scores. We identified the best-fit model for predicting WOMAC score using multiple MRI measures and patient-specific information, including sex, age, graft type, femoral transverse strain, femoral axial strain, and femoral shear strain. The best-fit model significantly predicted WOMAC score (p<0.001) better than any one individual MRI metric alone. When we regressed the model-predicted WOMAC scores against the patient-reported WOMAC scores, we found that our model achieved a goodness of fit exceeding 0.52. Conclusions: This work presents the first use of dualMRI in vivo in a cohort of participants at risk for developing osteoarthritis. Our results indicate that both shear and transverse strains are highly correlated with patient-reported outcome severity could serve as novel imaging biomarkers to predict the development of osteoarthritis.

Intervertebral Disc Elastography to Relate Shear Modulus and Relaxometry in Compression and Bending.

Intervertebral disc degeneration is the most recognized cause of low back pain, characterized by the decline of tissue structure and mechanics. Image-based mechanical parameters (e.g., strain, stiffness) may provide an ideal assessment of disc function that is lost with degeneration but unfortunately remains underdeveloped. Moreover, it is unknown whether strain or stiffness of the disc may be predicted by MRI relaxometry (e.g. T1 or T2), an increasingly accepted quantitative measure of disc structure. In this study, we quantified T1 and T2 relaxation times and in-plane strains using displacement-encoded MRI within the disc under physiological levels of compression and bending. We then estimated shear modulus in orthogonal image planes and compared these values to relaxation times and strains within regions of the disc. Intratissue strain depended on the loading mode, and shear modulus in the nucleus pulposus was typically an order of magnitude lower than the annulus fibrosis, except in bending, where the apparent stiffness depended on the loading. Relative shear moduli estimated from strain data derived under compression generally did not correspond with those from bending experiments, with no correlations in the sagittal plane and only 4 of 15 regions correlated in the coronal plane, suggesting that future inverse models should incorporate multiple loading conditions. Strain imaging and strain-based estimation of material properties may serve as imaging biomarkers to distinguish healthy and diseased discs. Additionally, image-based elastography and relaxometry may be viewed as complementary measures of disc structure and function to assess degeneration in longitudinal studies.

Demographic responses of hybridizing cinquefoils to changing climate in the Colorado Rocky Mountains.

Hybridization between taxa generates new pools of genetic variation that can lead to different environmental responses and demographic trajectories over time than seen in parental lineages. The potential for hybrids to have novel environmental tolerances may be increasingly important in mountainous regions, which are rapidly warming and drying due to climate change. Demographic analysis makes it possible to quantify within- and among-species responses to variation in climate and to predict population growth rates as those conditions change. We estimated vital rates and population growth in 13 natural populations of two cinquefoil taxa (Potentilla hippiana and P. pulcherrima) and their hybrid across elevation gradients in the Southern Rockies. Using three consecutive years of environmental and demographic data, we compared the demographic responses of hybrid and parental taxa to environmental variation across space and time. All three taxa had lower predicted population growth rates under warm, dry conditions. However, the magnitude of these responses varied among taxa and populations. Hybrids had consistently lower predicted population growth rates than P. hippiana. In contrast, hybrid performance relative to P. pulcherrima varied with population and climate, with the hybrid maintaining relatively stable growth rates while populations of P. pulcherrima shrank under warm, dry conditions. Our findings demonstrate that hybrids in this system are neither intrinsically unfit nor universally more vigorous than parents, suggesting that the demographic consequences of hybridization are context-dependent. Our results also imply that shifts to warmer and drier conditions could have particularly negative repercussions for P. pulcherrima, which is currently the most abundant taxon in the study area, possibly as a legacy of more favorable historical climates. More broadly, the distributions of these long-lived taxa are lagging behind their demographic trajectories, such that the currently less common P. hippiana could become the most abundant of the Potentilla taxa as this region continues to warm and dry.

Mechanical memory stored through epigenetic remodeling reduces cell therapeutic potential.

Understanding how cells remember previous mechanical environments to influence their fate, or mechanical memory, informs the design of biomaterials and therapies in medicine. Current regeneration therapies, such as cartilage regeneration procedures, require 2D cell expansion processes to achieve large cell populations critical for the repair of damaged tissues. However, the limit of mechanical priming for cartilage regeneration procedures before inducing long-term mechanical memory following expansion processes is unknown, and mechanisms defining how physical environments influence the therapeutic potential of cells remain poorly understood. Here, we identify a threshold to mechanical priming separating reversible and irreversible effects of mechanical memory. After 16 population doublings in 2D culture, expression levels of tissue-identifying genes in primary cartilage cells (chondrocytes) are not recovered when transferred to 3D hydrogels, while expression levels of these genes were recovered for cells only expanded for eight population doublings. Additionally, we show that the loss and recovery of the chondrocyte phenotype correlates with a change in chromatin architecture, as shown by structural remodeling of the trimethylation of H3K9. Efforts to disrupt the chromatin architecture by suppressing or increasing levels of H3K9me3 reveal that only with increased levels of H3K9me3 did the chromatin architecture of the native chondrocyte phenotype partially return, along with increased levels of chondrogenic gene expression. These results further support the connection between the chondrocyte phenotype and chromatin architecture, and also reveal the therapeutic potential of inhibitors of epigenetic modifiers as disruptors of mechanical memory when large numbers of phenotypically suitable cells are required for regeneration procedures.

Dynamic biophysical responses of neuronal cell nuclei and cytoskeletal structure following high impulse loading.

Cells are continuously exposed to dynamic environmental cues that influence their behavior. Mechanical cues can influence cellular and genomic architecture, gene expression, and intranuclear mechanics, providing evidence of mechanosensing by the nucleus, and a mechanoreciprocity between the nucleus and environment. Force disruption at the tissue level through aging, disease, or trauma, propagates to the nucleus and can have lasting consequences on proper functioning of the cell and nucleus. While the influence of mechanical cues leading to axonal damage has been well studied in neuronal cells, the mechanics of the nucleus following high impulse loading is still largely unexplored. Using an in vitro model of traumatic neural injury, we show a dynamic nuclear behavioral response to impulse stretch (up to 170% strain per second) through quantitative measures of nuclear movement, including tracking of rotation and internal motion. Differences in nuclear movement were observed between low and high strain magnitudes. Increased exposure to impulse stretch exaggerated the decrease in internal motion, assessed by particle tracking microrheology, and intranuclear displacements, assessed through high-resolution deformable image registration. An increase in F-actin puncta surrounding nuclei exposed to impulse stretch additionally demonstrated a corresponding disruption of the cytoskeletal network. Our results show direct biophysical nuclear responsiveness in neuronal cells through force propagation from the substrate to the nucleus. Understanding how mechanical forces perturb the morphological and behavioral response can lead to a greater understanding of how mechanical strain drives changes within the cell and nucleus, and may inform fundamental nuclear behavior after traumatic axonal injury. STATEMENT OF SIGNIFICANCE: The nucleus of the cell has been implicated as a mechano-sensitive organelle, courting molecular sensors and transmitting physical cues in order to maintain cellular and tissue homeostasis. Disruption of this network due to disease or high velocity forces (e.g., trauma) can not only result in orchestrated biochemical cascades, but also biophysical perturbations. Using an in vitro model of traumatic neural injury, we aimed to provide insight into the neuronal nuclear mechanics and biophysical responses at a continuum of strain magnitudes and after repetitive loads. Our image-based methods demonstrate mechanically-induced changes in cellular and nuclear behavior after high intensity loading and have the potential to further define mechanical thresholds of neuronal cell injury.

Spatial Gradients of Quantitative MRI as Biomarkers for Early Detection of Osteoarthritis: Data From Human Explants and the Osteoarthritis Initiative.

BACKGROUND: Healthy articular cartilage presents structural gradients defined by distinct zonal patterns through the thickness, which may be disrupted in the pathogenesis of several disorders. Analysis of textural patterns using quantitative MRI data may identify structural gradients of healthy or degenerating tissue that correlate with early osteoarthritis (OA). PURPOSE: To quantify spatial gradients and patterns in MRI data, and to probe new candidate biomarkers for early severity of OA. STUDY TYPE: Retrospective study. SUBJECTS: Fourteen volunteers receiving total knee replacement surgery (eight males/two females/four unknown, average age ± standard deviation: 68.1 ± 9.6 years) and 10 patients from the OA Initiative (OAI) with radiographic OA onset (two males/eight females, average age ± standard deviation: 57.7 ± 9.4 years; initial Kellgren-Lawrence [KL] grade: 0; final KL grade: 3 over the 10-year study). FIELD STRENGTH/SEQUENCE: 3.0-T and 14.1-T, biomechanics-based displacement-encoded imaging, fast spin echo, multi-slice multi-echo T2 mapping. ASSESSMENT: We studied structure and strain in cartilage explants from volunteers receiving total knee replacement, or structure in cartilage of OAI patients with progressive OA. We calculated spatial gradients of quantitative MRI measures (eg, T2) normal to the cartilage surface to enhance zonal variations. We compared gradient values against histologically OA severity, conventional relaxometry, and/or KL grades. STATISTICAL TESTS: Multiparametric linear regression for evaluation of the relationship between residuals of the mixed effects models and histologically determined OA severity scoring, with a significance threshold at α = 0.05. RESULTS: Gradients of individual relaxometry and biomechanics measures significantly correlated with OA severity, outperforming conventional relaxometry and strain metrics. In human explants, analysis of spatial gradients provided the strongest relationship to OA severity (R2  = 0.627). Spatial gradients of T2 from OAI data identified variations in radiographic (KL Grade 2) OA severity in single subjects, while conventional T2 alone did not. DATA CONCLUSION: Spatial gradients of quantitative MRI data may improve the predictive power of noninvasive imaging for early-stage degeneration. EVIDENCE LEVEL: 1 TECHNICAL EFFICACY: Stage 1.

Dispersal evolution in temporally variable environments: implications for plant range dynamics.

Dispersal-the movement of an individual from the site of birth to a different site for reproduction-is an ecological and evolutionary driver of species ranges that shapes patterns of colonization, connectivity, gene flow, and adaptation. In plants, the traits that influence dispersal often vary within and among species, are heritable, and evolve in response to the fitness consequences of moving through heterogeneous landscapes. Spatial and temporal variation in the quality and quantity of habitat are important sources of selection on dispersal strategies across species ranges. While recent reviews have evaluated the interactions between spatial variation in habitat and dispersal dynamics, the extent to which geographic variation in temporal variability can also shape range-wide patterns in dispersal traits has not been synthesized. In this paper, we summarize key predictions from metapopulation models that evaluate how dispersal evolves in response to spatial and temporal habitat variability. Next, we compile empirical data that quantify temporal variability in plant demography and patterns of dispersal trait variation across species ranges to evaluate the hypothesis that higher temporal variability favors increased dispersal at plant range limits. We found some suggestive evidence supporting this hypothesis while more generally identifying a major gap in empirical work evaluating plant metapopulation dynamics across species ranges and geographic variation in dispersal traits. To address this gap, we propose several future research directions that would advance our understanding of the interplay between spatiotemporal variability and dispersal trait variation in shaping the dynamics of current and future species ranges.

Draining the Swamping Hypothesis: Little Evidence that Gene Flow Reduces Fitness at Range Edges.

The genetic swamping hypothesis proposes that gene flow from central to peripheral populations inhibits local adaptation and is one of the most widely recognized explanations for range limitation. We evaluated empirical support for this hypothesis in studies quantifying patterns of gene flow to peripheral populations and their resulting fitness outcomes. We found little evidence that gene flow is generally asymmetric from central to peripheral populations and also that gene flow tends to have positive effects on edge population fitness. These findings contravene the long-held assumption that genetic swamping is a common driver of species range limits, and bear important implications for understanding the role of gene flow in range evolution and for predicting and managing eco-evolutionary responses to climate change.

The zonal evolution of collagen-network morphology quantified in early osteoarthritic grades of human cartilage.

Objective: We aimed to directly quantify the zone-specific evolution in morphology of collagen fibers and networks in human cartilage during the progression of early osteoarthritis. Collagen fibers exhibit depth-dependent orientations and diameters crucial to their mechanical roles. Cartilage degenerates in osteoarthritis, affecting the morphology of the collagen network and ultimately the intra-tissue mechanics. Design: We obtained specimens of human cartilage from healthy human knees ( n = 3 ) and from total knee arthroplasties ( n = 5 ). We utilized TEM and custom image analyses to visualize and quantify distributions in principal orientation, dispersion (about the principal orientation), and diameter of collagen fibers in the early grades of OA within each through-thickness zone. We then used histological and statistical analyses to probe for significant changes in the zone-specific evolution in collagen-network morphology as a function of Osteoarthritis Research Society International (OARSI) grade. Results: Dispersion in the alignment of collagen fibers increased with progression of early OA in both the superficial and deep zones, and decreased in the middle zone, while principal orientation did not change significantly. The non-normal and right-skewed distributions in fiber diameters did not evolve with the progression of OA. Conclusions: We provide the research community with quantitative data (1) on the through-thickness morphology of collagen in healthy cartilage and (2) on the evolution of through-thickness morphology of collagen with progressing early OA. Such quantitative data facilitate an improved mechanistic understanding of the progression of OA, and may facilitate identifying image-based biomarkers and treatment targets, and ultimately finding clinical interventions for OA.

Range edges in heterogeneous landscapes: Integrating geographic scale and climate complexity into range dynamics.

The impacts of climate change have re-energized interest in understanding the role of climate in setting species geographic range edges. Despite the strong focus on species' distributions in ecology and evolution, defining a species range edge is theoretically and empirically difficult. The challenge of determining a range edge and its relationship to climate is in part driven by the nested nature of geography and the multidimensionality of climate, which together generate complex patterns of both climate and biotic distributions across landscapes. Because range-limiting processes occur in both geographic and climate space, the relationship between these two spaces plays a critical role in setting range limits. With both conceptual and empirical support, we argue that three factors-climate heterogeneity, collinearity among climate variables, and spatial scale-interact to shape the spatial structure of range edges along climate gradients, and we discuss several ways that these factors influence the stability of species range edges with a changing climate. We demonstrate that geographic and climate edges are often not concordant across species ranges. Furthermore, high climate heterogeneity and low climate collinearity across landscapes increase the spectrum of possible relationships between geographic and climatic space, suggesting that geographic range edges and climatic niche limits correspond less frequently than we may expect. More empirical explorations of how the complexity of real landscapes shapes the ecological and evolutionary processes that determine species range edges will advance the development of range limit theory and its applications to biodiversity conservation in the context of changing climate.

The Effects of Temporal Variation on Fitness, Functional Traits, and Species Distribution Patterns.

Temporal variation is a powerful source of selection on life history strategies and functional traits in natural populations. Theory predicts that the rate and predictability of fluctuations should favor distinct strategies, ranging from phenotypic plasticity to bet-hedging, which are likely to have important consequences for species distribution patterns and their responses to environmental change. To date, we have few empirical studies that test those predictions in natural systems, and little is known about how genetic, environmental, and developmental factors interact to define the "fluctuation niche" of species in temporally variable environments. In this study, we evaluated the effects of hydrological variability on fitness and functional trait variation in three closely related plant species in the genus Lasthenia that occupy different microhabitats within vernal pool landscapes. Using a controlled greenhouse experiment, we manipulated the mean and variability in hydrological conditions by growing plants at different depths with respect to a shared water table and manipulating the magnitude of stochastic fluctuations in the water table over time. We found that all species had similarly high relative fitness above the water table, but differed in their sensitivities to water table fluctuations. Specifically, the two species from vernal pools basins, where soil moisture is controlled by a perched water table, were negatively affected by the stochasticity treatments. In contrast, a species from the upland habitat surrounding vernal pools, where stochastic precipitation events control soil moisture variation, was insensitive to experimental fluctuations in the water table. We found strong signatures of genetic, environmental (plastic), and developmental variation in four traits that can influence plant hydrological responses. Three of these traits varied across plant development and among experimental treatments in directions that aligned with constitutive differences among species, suggesting that multiple sources of variation align to facilitate phenotypic matching with the hydrological environment in Lasthenia. We found little evidence for predicted patterns of phenotypic plasticity and bet-hedging in species and traits from predictable and stochastic environments, respectively. We propose that selection for developmental shifts in the hydrological traits of Lasthenia species has reduced or modified selection for plasticity at any given stage of development. Collectively, these results suggest that variation in species' sensitivities to hydrological stochasticity may explain why vernal pool Lasthenia species do not occur in upland habitat, and that all three species integrate genetic, environmental, and developmental information to manage the unique patterns of temporal hydrological variation in their respective microhabitats.

Grow Where You Thrive, or Where Only You Can Survive? An Analysis of Performance Curve Evolution in a Clade with Diverse Habitat Affinities.

Performance curves are valuable tools for quantifying the fundamental niches of organisms and testing hypotheses about evolution, life-history trade-offs, and the drivers of variation in species' distribution patterns. Here, we present a novel Bayesian method for characterizing performance curves that facilitates comparisons among species. We then use this model to quantify and compare the hydrological performance curves of 14 different taxa in the genus Lasthenia, an ecologically diverse clade of plants that collectively occupy a variety of habitats with unique hydrological features, including seasonally flooded wetlands called vernal pools. We conducted a growth chamber experiment to measure each taxon's fitness across five hydrological treatments that ranged from severe drought to extended flooding, and we identified differences in hydrological performance curves that explain their associations with vernal pool and terrestrial habitats. Our analysis revealed that the distribution of vernal pool taxa in the field does not reflect their optimal hydrological environments: all taxa, regardless of habitat affinity, have highest fitness under similar hydrological conditions of saturated soil without submergence. We also found that a taxon's relative position across flood gradients within vernal pools is best predicted by the height of its performance curve. These results demonstrate the utility of our approach for generating insights into when and how performance curves evolve among taxa as they diversify into distinct environments. To facilitate its use, the modeling framework has been developed into an R package.

The adaptive potential of plant populations in response to extreme climate events.

The frequency and magnitude of extreme climate events are increasing with global change, yet we lack predictions and empirical evidence for the ability of wild populations to persist and adapt in response to these events. Here, we used Fisher's Fundamental Theorem of Natural Selection to evaluate the adaptive potential of Lasthenia fremontii, a herbaceous winter annual that is endemic to seasonally flooded wetlands in California, to alternative flooding regimes that occur during El Niño Southern Oscillation (ENSO) events. The results indicate that populations may exhibit greater adaptive potential in response to dry years than wet years, and that the relative performance of populations will change across climate scenarios. More generally, our findings show that extreme climate events can substantially change the potential for populations to adapt to climate change by modulating the expression of standing genetic variation and mean fitness.

Conserved thermal performance curves across the geographic range of a gametophytic fern.

Species-level responses to environmental change depend on the collective responses of their constituent populations and the degree to which populations are specialized to local conditions. Manipulative experiments in common-garden settings make it possible to test for population variation in species' responses to specific climate variables, including those projected to shift as the climate changes in the future. While this approach is being applied to a variety of plant taxa to evaluate their responses to climate change, these studies are heavily biased towards seed-bearing plant species. Given several unique morphological and physiological traits, fern species may exhibit very different responses from angiosperms and gymnosperms. Here, we tested the hypothesis that previously detected population differentiation in a fern species is due to differentiation in thermal performance curves among populations. We collected explants from six populations spanning the species' geographic range and exposed them to 10 temperature treatments. Explant survival, lifespan and the change in photosynthetic area were analysed as a function of temperature, source population and their interaction. Overall results indicated that explants performed better at the lowest temperature examined, and the threshold for explant performance reflects maximum temperatures likely to be experienced in the field. Surprisingly, explant fitness did not differ among source populations, suggesting that temperature is not the driver behind previously detected patterns of population differentiation. These results highlight the importance of other environmental axes in driving population differentiation across a species range, and suggest that the perennial life history strategy, asexual mating system and limited dispersal potential of Vittaria appalachiana may restrict the rise and differentiation of adaptive genetic variation in thermal performance traits among populations.

Environmental predictors of dispersal traits across a species' geographic range.

Variation in habitat quality and quantity drive selection on dispersal traits in heterogeneous environments, but the extent to which environmental conditions predict geographic variation in dispersal is rarely evaluated. We assessed dispersal trait variation across the range of Cakile edentula var. lacustris, an annual herb that occupies beaches of the Great Lakes. Cakile edentula has dimorphic fruits that each contain one dispersive and one non-dispersive seed. Previous work showed that plant height, branching density, and dispersive fruit wing-loading can determine the distance that seeds disperse locally by wind, while pericarp thickness influences the distance they disperse by water. We tested if these traits vary predictably with latitude across the species' geographic range, and if variation in dispersal characteristics can be predicted by the quality and quantity of habitat available at a site. We observed that the dispersive fruits from northern and southern populations had thinner pericarps than those from the interior of the species' range, reflecting reduced long-distance dispersal by water at both range limits. Plants at the northern range limit were shorter with less dense branching and lower wing-loading than populations elsewhere in the range, suggesting that these populations have enhanced local wind dispersal. In contrast, southern populations exhibited traits with inconsistent effects on wind dispersal: plants tended to be short, which facilitates wind dispersal in C. edentula, but also had relatively higher branching density and distal segment wing-loading that reduce wind dispersal. Geographic variation in maternal plant height and branching density was partially explained by variation in habitat quality, which declined at the species' range limits. In addition, population differences in branching density, fruit wing-loading, and pericarp thickness were predicted by the abundance and distribution of beach habitat. Finally, a common garden analysis recovered latitudinal patterns for the dispersal traits associated with fruits, but not those associated with maternal architecture. Thus, the geographic patterns of dispersal trait variation that we observed likely reflect responses to past selection by the distribution, abundance, and quality of habitat, strong plasticity in dispersal traits, and the effects dispersal itself has in shaping local adaptation by driving gene flow among populations.

Facilitative and competitive interaction components among New England salt marsh plants.

Intra- and interspecific interactions can be broken down into facilitative and competitive components. The net interaction between two organisms is simply the sum of these counteracting elements. Disentangling the positive and negative components of species interactions is a critical step in advancing our understanding of how the interaction between organisms shift along physical and biotic gradients. We performed a manipulative field experiment to quantify the positive and negative components of the interactions between a perennial forb, Aster tenuifolius, and three dominant, matrix-forming grasses and rushes in a New England salt marsh. Specifically, we asked whether positive and negative interaction components: (1) are unique or redundant across three matrix-forming species (two grasses; Distichlis spicata and Spartina patens, and one rush; Juncus gerardi), and (2) change across Aster life stages (seedling, juvenile, and adult). For adult Aster the strength of the facilitative component of the matrix-forb interaction was stronger than the competitive component for two of the three matrix species, leading to net positive interactions. There was no statistically significant variation among matrix species in their net or component effects. We found little difference in the effects of J. gerardi on Aster at later life-history stages; interaction component strengths did not differ between juveniles and adults. However, mortality of seedlings in neighbor removal plots was 100%, indicating a particularly strong and critical facilitative effect of matrix species on this forb during the earliest life stages. Overall, our results indicate that matrix forming grasses and rushes have important, yet largely redundant, positive net effects on Aster performance across its life cycle. Studies that untangle various components of interactions and their contingencies are critical to both expanding our basic understanding of community organization, and predicting how natural communities and their component parts will respond to environmental change.

Habitat filtering not dispersal limitation shapes oceanic island floras: species assembly of the Galápagos archipelago.

Remote locations, such as oceanic islands, typically harbour relatively few species, some of which go on to generate endemic radiations. Species colonising these locations tend to be a non-random subset from source communities, which is thought to reflect dispersal limitation. However, non-random colonisation could also result from habitat filtering, whereby only a few continental species can become established. We evaluate the imprints of these processes on the Galápagos flora by analysing a comprehensive regional phylogeny for ~ 39 000 species alongside information on dispersal strategies and climatic suitability. We found that habitat filtering was more important than dispersal limitation in determining species composition. This finding may help explain why adaptive radiation is common on oceanic archipelagoes - because colonising species can be relatively poor dispersers with specific niche requirements. We suggest that the standard assumption that plant communities in remote locations are primarily shaped by dispersal limitation deserves reconsideration.

Cyclic loading of human articular cartilage: The transition from compaction to fatigue.

Osteoarthritis and articular cartilage injuries are common conditions in human joints and a frequent cause of pain and disability. Unfortunately, cartilage is avascular and has limited capabilities for self-repair. Despite the societal impact, there is little information on the dynamic process of cartilage degeneration. We performed a series of cyclic unconfined compression tests motivated by in vivo loading conditions and designed to generate mechanical fatigue. We examined the functional (both stress-stretch and creep) responses of the tissue after recovery from a specified number of loading cycles, as well as histology and second harmonic generation microscopy images. The effect of compaction was complimented by the effect of fatigue in our unconfined compression tests. A three-way, repeated-measures mixed model ANOVA showed significant differences between loading with a physiologically relevant low magnitude, and two more severe loading magnitudes, in terms of the resulting specimen stiffness, time to equilibrium and thickness. There was a statistically significant effect of loading frequency on a specimen's time to equilibrium and significant interaction of force and frequency on specimen thickness and time to equilibrium. Increasing the number of loading cycles significantly impacted a specimen's effective stiffness and resulting thickness. We attribute permanent loss of mechanical function under cyclic loading to rearrangement and disruption of the collagen network and resulting proteoglycan (PG) aggregation, as seen in histological and second harmonic generation images, as a result of induced mechanical fatigue.

Spatiotemporal heterogeneity in precipitation patterns explain population-level germination strategies in an edaphic specialist.

BACKGROUND AND AIMS: Many locally endemic species in biodiversity hotspots are restricted to edaphic conditions that are fixed in the landscape, limiting their potential to track climate change through dispersal. Instead, such species experience strong selection for germination strategies that can track suitable conditions through time. Germination strategies were compared among populations across the geographic range of a California vernal pool annual, Lasthenia fremontii Local germination strategies were tested to determine the associations with geographic variation in precipitation patterns. METHODS: This study evaluated patterns of seed germination, dormancy and mortality in response to simulated variation in the timing, amount and duration of the first autumn precipitation event using seeds from six populations that span a geographic gradient in precipitation. Next, it was tested whether the germination strategies of different populations can be predicted by historical precipitation patterns that characterize each site. KEY RESULTS: A significant positive relationship was observed between the historical variability in autumn precipitation and the extent of dormancy in a population. Marginal populations, with histories of the most extreme but constant autumn precipitation levels, expressed the lowest dormancy levels. Populations from sites with historically higher levels of autumn precipitation tended to germinate faster, but this tendency was not statistically significant. CONCLUSIONS: Germination in L. fremontii is cued by the onset of the first rains that characterize the beginning of winter in California's Great Central Valley. However, populations differ in how fast they germinate and the fraction of seeds that remain dormant when germination cues occur. The results suggest that seed dormancy may be a key trait for populations to track increasingly drier climates predicted by climate change models. However, the low dormancy and high mortality levels observed among seeds of the southernmost, driest populations make them most vulnerable to local extinction.