Belowground plant parts are crucial for comprehensively estimating total plant richness in herbaceous and woody habitats.

Most studies consider aboveground plant species richness as a representative biodiversity measure. This approach inevitably assumes that the partitioning of total plant species richness into above- and belowground components is constant or at least consistent within and across vegetation types. However, with studies considering belowground plant richness still scarce and completely absent along vegetation gradients, this assumption lacks experimental support. Novel DNA sequencing techniques allow economical, high-throughput species identification of belowground environmental samples, enabling the measurement of the contributions of both above- and belowground plant components to total plant richness. We investigated above- and belowground plant species richness in four vegetation types (birch forest, heath, low alpine tundra, high alpine tundra) at the scale of herbaceous plant neighborhoods (dm) using 454 sequencing of the chloroplast trnL (UAA) intron to determine the plant species richness of environmental root samples and combined it with aboveground data from vegetation surveys to obtain total plant species richness. We correlated the measured plant species richness components with each other and with their respective plant biomass components within and across vegetation types. Total plant species richness exceeded aboveground richness twice on average and by as much as three times in low alpine tundra, indicating that a significant fraction of belowground plant richness cannot be recorded aboveground. More importantly, no consistent relationship among richness components (above- and belowground) was found within or across vegetation types, indicating that aboveground richness alone cannot predict total plant richness in contrasting vegetation types. Finally, no consistent relationship between plant richness and the corresponding biomass component was found. Our results clearly show that aboveground plant richness alone is a poor estimator of total plant species richness within and across different vegetation types. Consequently, it is crucial to account for belowground plant richness in future plant ecological studies in order to validate currently accepted plant richness patterns, as well as to measure potential changes in plant community composition in a changing environment.

Preventing vessel perforations in endovascular thrombectomy: feasibility and safety of passing the clot with a microcatheter without microwire: the wireless microcatheter technique.

BACKGROUND: To place a stent retriever for thrombectomy in acute ischemic stroke, the clot has to be passed first. A microwire is usually used for this maneuver. As an alternative, a wireless microcatheter can be used to pass the clot. OBJECTIVE: To analyze the feasibility and complication rates of passing the clot using either a microwire or a wireless microcatheter. METHODS: A retrospective non-randomized analysis of 110 consecutive patients with acute ischemic stroke in the anterior circulation was performed, in whom video recordings of mechanical thrombectomies were available. In total, 203 attempts at mechanical recanalization were performed. RESULTS: Successful recanalization (TICI 2b-3) was achieved in 97.3% of patients. In 71.8% of attempts the clot was successfully passed using a wireless microcatheter only. When a microwire was used initially, clot passage was successful in 95.3% of attempts. Complication rates for angiographically detectable subarachnoid hemorrhage were 6.1% when a microwire was used to pass the clot compared with 0% when a wireless microcatheter was used (p<0.001). Complication rates for angiographically occult circumscribed subarachnoid contrast extravasation observed on post-interventional CT scans were 18.2% when a microwire was used to pass the clot and 4.5% when a wireless microcatheter was used (p<0.001). CONCLUSIONS: In most cases of mechanical recanalization the clot can be passed with a wireless microcatheter instead of a microwire. In our study this method significantly reduced the risk for vessel perforation and subarachnoid hemorrhage. We therefore recommend the use of this technique whenever possible.

Future soil moisture and temperature extremes imply expanding suitability for rainfed agriculture in temperate drylands.

The distribution of rainfed agriculture, which accounts for approximately ¾ of global croplands, is expected to respond to climate change and human population growth and these responses may be especially pronounced in water limited areas. Because the environmental conditions that support rainfed agriculture are determined by climate, weather, and soil conditions that affect overall and transient water availability, predicting this response has proven difficult, especially in temperate regions that support much of the world's agriculture. Here, we show that suitability to support rainfed agriculture in temperate dryland climates can be effectively represented by just two daily environmental variables: moist soils with warm conditions increase suitability while extreme high temperatures decrease suitability. 21st century projections based on daily ecohydrological modeling of downscaled climate forecasts indicate overall increases in the area suitable for rainfed agriculture in temperate dryland regions, especially at high latitudes. The regional exception to this trend was Europe, where suitability in temperate dryland portions will decline substantially. These results clarify how rising temperatures interact with other key drivers of moisture availability to determine the sustainability of rainfed agriculture and help policymakers, resource managers, and the agriculture industry anticipate shifts in areas suitable for rainfed cultivation.

Climate change reduces extent of temperate drylands and intensifies drought in deep soils.

Drylands cover 40% of the global terrestrial surface and provide important ecosystem services. While drylands as a whole are expected to increase in extent and aridity in coming decades, temperature and precipitation forecasts vary by latitude and geographic region suggesting different trajectories for tropical, subtropical, and temperate drylands. Uncertainty in the future of tropical and subtropical drylands is well constrained, whereas soil moisture and ecological droughts, which drive vegetation productivity and composition, remain poorly understood in temperate drylands. Here we show that, over the twenty first century, temperate drylands may contract by a third, primarily converting to subtropical drylands, and that deep soil layers could be increasingly dry during the growing season. These changes imply major shifts in vegetation and ecosystem service delivery. Our results illustrate the importance of appropriate drought measures and, as a global study that focuses on temperate drylands, highlight a distinct fate for these highly populated areas.

Potential contributions of root decomposition to the nitrogen cycle in arctic forest and tundra.

Plant contributions to the nitrogen (N) cycle from decomposition are likely to be altered by vegetation shifts associated with climate change. Roots account for the majority of soil organic matter input from vegetation, but little is known about differences between vegetation types in their root contributions to nutrient cycling. Here, we examine the potential contribution of fine roots to the N cycle in forest and tundra to gain insight into belowground consequences of the widely observed increase in woody vegetation that accompanies climate change in the Arctic. We combined measurements of root production from minirhizotron images with tissue analysis of roots from differing root diameter and color classes to obtain potential N input following decomposition. In addition, we tested for changes in N concentration of roots during early stages of decomposition, and investigated whether vegetation type (forest or tundra) affected changes in tissue N concentration during decomposition. For completeness, we also present respective measurements of leaves. The potential N input from roots was twofold greater in forest than in tundra, mainly due to greater root production in forest. Potential N input varied with root diameter and color, but this variation tended to be similar in forest and tundra. As for roots, the potential N input from leaves was significantly greater in forest than in tundra. Vegetation type had no effect on changes in root or leaf N concentration after 1 year of decomposition. Our results suggest that shifts in vegetation that accompany climate change in the Arctic will likely increase plant-associated potential N input both belowground and aboveground. In contrast, shifts in vegetation might not alter changes in tissue N concentration during early stages of decomposition. Overall, differences between forest and tundra in potential contribution of decomposing roots to the N cycle reinforce differences between habitats that occur for leaves.

Climate change-induced vegetation shifts lead to more ecological droughts despite projected rainfall increases in many global temperate drylands.

Drylands occur worldwide and are particularly vulnerable to climate change because dryland ecosystems depend directly on soil water availability that may become increasingly limited as temperatures rise. Climate change will both directly impact soil water availability and change plant biomass, with resulting indirect feedbacks on soil moisture. Thus, the net impact of direct and indirect climate change effects on soil moisture requires better understanding. We used the ecohydrological simulation model SOILWAT at sites from temperate dryland ecosystems around the globe to disentangle the contributions of direct climate change effects and of additional indirect, climate change-induced changes in vegetation on soil water availability. We simulated current and future climate conditions projected by 16 GCMs under RCP 4.5 and RCP 8.5 for the end of the century. We determined shifts in water availability due to climate change alone and due to combined changes of climate and the growth form and biomass of vegetation. Vegetation change will mostly exacerbate low soil water availability in regions already expected to suffer from negative direct impacts of climate change (with the two RCP scenarios giving us qualitatively similar effects). By contrast, in regions that will likely experience increased water availability due to climate change alone, vegetation changes will counteract these increases due to increased water losses by interception. In only a small minority of locations, climate change-induced vegetation changes may lead to a net increase in water availability. These results suggest that changes in vegetation in response to climate change may exacerbate drought conditions and may dampen the effects of increased precipitation, that is, leading to more ecological droughts despite higher precipitation in some regions. Our results underscore the value of considering indirect effects of climate change on vegetation when assessing future soil moisture conditions in water-limited ecosystems.

New Multicentury Evidence for Dispersal Limitation during Primary Succession.

Primary succession is limited by both ecosystem development and plant dispersal, but the extent to which dispersal constrains succession over the long-term is unknown. We compared primary succession along two co-occurring arctic chronosequences with contrasting spatial scales: sorted circles that span a few meters and may have few dispersal constraints and glacial forelands that span several kilometers and may have greater dispersal constraints. Dispersal constraints slowed primary succession by centuries: plots were dominated by cryptogams after 20 years on circles but after 270 years on forelands; plots supported deciduous plants after 100 years on circles but after >400 years on forelands. Our study provides century-scale evidence suggesting that dispersal limitations constrain the rate of primary succession in glacial forelands.

The hidden season: growing season is 50% longer below than above ground along an arctic elevation gradient.

There is compelling evidence from experiments and observations that climate warming prolongs the growing season in arctic regions. Until now, the start, peak, and end of the growing season, which are used to model influences of vegetation on biogeochemical cycles, were commonly quantified using above-ground phenological data. Yet, over 80% of the plant biomass in arctic regions can be below ground, and the timing of root growth affects biogeochemical processes by influencing plant water and nutrient uptake, soil carbon input and microbial activity. We measured timing of above- and below-ground production in three plant communities along an arctic elevation gradient over two growing seasons. Below-ground production peaked later in the season and was more temporally uniform than above-ground production. Most importantly, the growing season continued c. 50% longer below than above ground. Our results strongly suggest that traditional above-ground estimates of phenology in arctic regions, including remotely sensed information, are not as complete a representation of whole-plant production intensity or duration, as studies that include root phenology. We therefore argue for explicit consideration of root phenology in studies of carbon and nutrient cycling, in terrestrial biosphere models, and scenarios of how arctic ecosystems will respond to climate warming.

Species richness of arbuscular mycorrhizal fungi: associations with grassland plant richness and biomass.

Although experiments show a positive association between vascular plant and arbuscular mycorrhizal fungal (AMF) species richness, evidence from natural ecosystems is scarce. Furthermore, there is little knowledge about how AMF richness varies with belowground plant richness and biomass. We examined relationships among AMF richness, above- and belowground plant richness, and plant root and shoot biomass in a native North American grassland. Root-colonizing AMF richness and belowground plant richness were detected from the same bulk root samples by 454-sequencing of the AMF SSU rRNA and plant trnL genes. In total we detected 63 AMF taxa. Plant richness was 1.5 times greater belowground than aboveground. AMF richness was significantly positively correlated with plant species richness, and more strongly with below- than aboveground plant richness. Belowground plant richness was positively correlated with belowground plant biomass and total plant biomass, whereas aboveground plant richness was positively correlated only with belowground plant biomass. By contrast, AMF richness was negatively correlated with belowground and total plant biomass. Our results indicate that AMF richness and plant belowground richness are more strongly related with each other and with plant community biomass than with the plant aboveground richness measures that have been almost exclusively considered to date.

Reduction in distal emboli with proximal flow control during mechanical thrombectomy: a quantitative in vitro study.

BACKGROUND AND PURPOSE: To evaluate the impact of proximal flow control on efficacy and safety of mechanical thrombectomy in an in vitro middle cerebral artery occlusion. METHODS: Three independent variables, including clot type, device (Merci Retriever, Solitaire FR, and Trevo devices), and use of a balloon guide catheter, were used to ascertain the impact of proximal flow control on the size and number of distal emboli generated during thrombectomy. Secondary end points were the recanalization rate and amount of flow restored. RESULTS: Use of the balloon guide catheter during thrombectomy of the fragile, hard clot significantly reduced the formation of large distal emboli with a diameter >1 mm, regardless of the device used (P<0.01). Applying aspiration via the balloon guide catheter in place of the conventional guide catheter resulted in a significant increase of flow reversal (P<0.0001). Prior to thrombectomy, deployment of the stent-trievers produced immediate flow restoration through the soft and hard clot occlusions, 69.2 ± 27.3 and 45.5 ± 22.8 mL/min, respectively, that was preserved after the balloon inflation because of collateral flow via the posterior communication artery. After deployment but before thrombectomy, no flow was restored when using the Merci Retriever. After thrombectomy, complete flow restoration was achieved in a majority of cases. The Merci Retriever required more thrombectomy attempts to achieve hard clot removal compared with the stent-trievers when the conventional guide catheter was used (1.5 versus 1.1). CONCLUSIONS: The risk of distal embolization was significantly reduced with the use of the balloon guide catheter.

Plant species richness belowground: higher richness and new patterns revealed by next-generation sequencing.

Variation in plant species richness has been described using only aboveground vegetation. The species richness of roots and rhizomes has never been compared with aboveground richness in natural plant communities. We made direct comparisons of grassland plant richness in identical volumes (0.1 × 0.1 × 0.1 m) above and below the soil surface, using conventional species identification to measure aboveground richness and 454 sequencing of the chloroplast trnL(UAA) intron to measure belowground richness. We described above- and belowground richness at multiple spatial scales (from a neighbourhood scale of centimetres to a community scale of hundreds of metres), and related variation in richness to soil fertility. Tests using reference material indicated that 454 sequencing captured patterns of species composition and abundance with acceptable accuracy. At neighbourhood scales, belowground richness was up to two times greater than aboveground richness. The relationship between above- and belowground richness was significantly different from linear: beyond a certain level of belowground richness, aboveground richness did not increase further. Belowground richness also exceeded that of aboveground at the community scale, indicating that some species are temporarily dormant and absent aboveground. Similar to other grassland studies, aboveground richness declined with increasing soil fertility; in contrast, the number of species found only belowground increased significantly with fertility. These results indicate that conventional aboveground studies of plant richness may overlook many coexisting species, and that belowground richness becomes relatively more important in conditions where aboveground richness decreases. Measuring plant belowground richness can considerably alter perceptions of biodiversity and its responses to natural and anthropogenic factors.

The invasive grass Agropyron cristatum doubles belowground productivity but not soil carbon.

Root dynamics are among the largest knowledge gaps in determining how terrestrial carbon (C) cycles will respond to environmental change. Increases in productivity accompanying plant invasions and introductions could increase ecosystem C storage, but belowground changes are unknown, even though roots may account for 50-90% of production in temperate ecosystems. We examined whether the introduction of a widespread invasive grass with relatively high shoot production also increased belowground productivity and soil C storage, using a multiyear rhizotron study in 50-year-old stands dominated either by the invasive C3 grass Agropyron cristatum or by largely C4 native grasses. Relative to native vegetation, stands dominated by the invader had doubled root productivity. Soil carbon isotope values showed that the invader had made detectable contributions to soil C. Soil C content, however, was not significantly different between invader-dominated stands (0.42 mg C/g soil) and native vegetation (0.45 mg C/g soil). The discrepancy between enhanced production and lack of soil C changes was attributable to differences in root traits between invader-dominated stands and native vegetation. Relative to native vegetation, roots beneath the invader had 59% more young white tissue, with 80% higher mortality and 19% lower C:N ratios (all P < 0.05). Such patterns have previously been reported for aboveground tissues of invaders, and we show that they are also found belowground. If these root traits occur in other invasive species, then the global phenomenon of increased productivity following biological invasion may not increase soil C storage.

Herbivory limits recruitment in an old-field seed addition experiment.

Environmental variability can promote coexistence by creating establishment sites for rare plants, but low diversity in anthropogenic grasslands suggests that this variability may be eliminated (homogenization hypothesis) or inaccessible (barrier hypothesis). We explore these alternatives on the northern Great Plains, where 11 million hectares have been transformed by multiple environmental changes, but the causes of species loss are unclear. In a degraded grassland, we increased environmental variability by manipulating competition and herbivory along gradients of fertility and disturbance, and we circumvented dispersal barriers by adding 1.2 million seeds of five functionally distinct species at varying densities. The experiment ended after 12 weeks due to the direct and indirect effects of unapparent small native herbivores, which were barriers to population establishment by the added species. The direct cause of recruitment failure was browsing. The indirect cause was associated with competition from invasive plants that appeared to be more tolerant or resistant to herbivory. Variability in fertility, disturbance, propagule pressure, and competition had relatively minor impacts on colonization by the added species because herbivores controlled recruitment in most environments. Recruitment outside the herbivore exclosures was mostly by unpalatable exotics, suggesting a possible link between invasion success and herbivore resistance for some introduced plants.

Competition, resources, and vegetation during 10 years in native grassland.

A 10-year experiment tested for variation in competition intensity over time in a natural grassland at the northern edge of the Great Plains. Growing-season precipitation varied fivefold during the study. All ecosystem-level variables varied significantly among years, and most covaried in expected ways. The covers of all common grasses possessing the C3 photosynthetic pathway varied significantly among years; in contrast, all common species with traits associated with drought tolerance (a C4 grass, a lichen, a spikemoss, and a subshrub) did not vary. Annual transplant experiments measured the competitive effects of neighbors on the growth of individuals of the native grass Bouteloua gracilis. A significant interaction between year and competition showed that competition intensity varied among years. The size of this effect, however, was small (eta2 = 0.074) relative to the size of the direct effect of competition (eta2 = 0.20) or the year in which the experiment was conducted (eta2 = 0.51). Further, competition intensity was not significantly related to any variable describing standing crop or resources, or species richness. Species richness was highest in years with high precipitation, standing crop, and individual growth, due to the recruitment of rare species that were absent from dry years. In summary, variation in competition intensity was statistically significant but had small effects relative to the direct effects of climate.

Interactive effects of fertilization and disturbance on community structure and resource availability in an old-field plant community.

The interactive effects of fertilization and disturbance on plant community structure and resource availability were studied by supplying four levels of nitrogen and applying four intensities of tilling to a 30 year old field in a factorial design for 2 year. Live above-ground biomass, root biomass, and litter generally increased with nitrogen supply and decreased with disturbance. Species composition varied significantly, with annuals increasing with both nitrogen and disturbance, but with perennials unaffected by nitrogen and decreased by disturbance. Species diversity decreased with disturbance, but decreased with nitrogen only in undisturbed vegetation. Root: shoot ratios decreased with added nitrogen, leaf allocation decreased with disturbance, and flowering allocation increased. Surprisingly, stem allocation was unaffected by disturbance. This result reflected a shift from vertical stems to horizontal stems as disturbance increased. Resource measurements suggested that the vegetation responded to interactions between the treatments as well as to direct treatment effects. Variation in light penetration was reduced by fertilization in undisturbed vegetation but not in tilled plots; variability was not directly affected by disturbance. The availability of nitrogen, the limiting soil nutrient, increased with fertilization but was not significantly affected by disturbance. In contrast, the ratio of ammonium to nitrate was significantly reduced by disturbance but unaffected by supply rates, suggesting that nitrogen may have had different effects under different disturbance regimes, even though its total availability was constant. While many community responses to fertilization and disturbance conformed to those reported earlier, resource and allocation measurements indicated that their interactions are not always predictable from their separate effects.