Drivers of soil microbial and detritivore activity across global grasslands.

Covering approximately 40% of land surfaces, grasslands provide critical ecosystem services that rely on soil organisms. However, the global determinants of soil biodiversity and functioning remain underexplored. In this study, we investigate the drivers of soil microbial and detritivore activity in grasslands across a wide range of climatic conditions on five continents. We apply standardized treatments of nutrient addition and herbivore reduction, allowing us to disentangle the regional and local drivers of soil organism activity. We use structural equation modeling to assess the direct and indirect effects of local and regional drivers on soil biological activities. Microbial and detritivore activities are positively correlated across global grasslands. These correlations are shaped more by global climatic factors than by local treatments, with annual precipitation and soil water content explaining the majority of the variation. Nutrient addition tends to reduce microbial activity by enhancing plant growth, while herbivore reduction typically increases microbial and detritivore activity through increased soil moisture. Our findings emphasize soil moisture as a key driver of soil biological activity, highlighting the potential impacts of climate change, altered grazing pressure, and eutrophication on nutrient cycling and decomposition within grassland ecosystems.

Fire shapes fungal guild diversity and composition through direct and indirect pathways.

Fire has shaped global ecosystems for millennia by directly killing organisms and indirectly altering habitats and resources. All terrestrial ecosystems, including fire-prone ecosystems, rely on soil-inhabiting fungi, where they play vital roles in ecological processes. Yet our understanding of how fire regimes influence soil fungi remains limited and our knowledge of these interactions in semiarid landscapes is virtually absent. We collected soil samples and vegetation measurements from sites across a gradient in time-since-fire ages (0-75 years-since-fire) and fire frequency (burnt 0-5 times during the recent 29-year period) in a semiarid heathland of south-eastern Australia. We characterized fungal communities using ITS amplicon-sequencing and assigned fungi taxonomically to trophic guilds. We used structural equation models to examine direct, indirect and total effects of time-since-fire and fire frequency on total fungal, ectomycorrhizal, saprotrophic and pathogenic richness. We used multivariate analyses to investigate how total fungal, ectomycorrhizal, saprotrophic and pathogenic species composition differed between post-fire successional stages and fire frequency classes. Time-since-fire was an important driver of saprotrophic richness; directly, saprotrophic richness increased with time-since-fire, and indirectly, saprotrophic richness declined with time-since-fire (resulting in a positive total effect), mediated through the impact of fire on substrates. Frequently burnt sites had lower numbers of saprotrophic and pathogenic species. Post-fire successional stages and fire frequency classes were characterized by distinct fungal communities, with large differences in ectomycorrhizal species composition. Understanding the complex responses of fungal communities to fire can be improved by exploring how the effects of fire flow through ecosystems. Diverse fire histories may be important for maintaining the functional diversity of fungi in semiarid regions.

Evolutionary history of grazing and resources determine herbivore exclusion effects on plant diversity.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory.

Temporal rarity is a better predictor of local extinction risk than spatial rarity.

Spatial rarity is often used to predict extinction risk, but rarity can also occur temporally. Perhaps more relevant in the context of global change is whether a species is core to a community (persistent) or transient (intermittently present), with transient species often susceptible to human activities that reduce niche space. Using 5-12 yr of data on 1,447 plant species from 49 grasslands on five continents, we show that local abundance and species persistence under ambient conditions are both effective predictors of local extinction risk following experimental exclusion of grazers or addition of nutrients; persistence was a more powerful predictor than local abundance. While perturbations increased the risk of exclusion for low persistence and abundance species, transient but abundant species were also highly likely to be excluded from a perturbed plot relative to ambient conditions. Moreover, low persistence and low abundance species that were not excluded from perturbed plots tended to have a modest increase in abundance following perturbance. Last, even core species with high abundances had large decreases in persistence and increased losses in perturbed plots, threatening the long-term stability of these grasslands. Our results demonstrate that expanding the concept of rarity to include temporal dynamics, in addition to local abundance, more effectively predicts extinction risk in response to environmental change than either rarity axis predicts alone.

Negative effects of nitrogen override positive effects of phosphorus on grassland legumes worldwide.

Anthropogenic nutrient enrichment is driving global biodiversity decline and modifying ecosystem functions. Theory suggests that plant functional types that fix atmospheric nitrogen have a competitive advantage in nitrogen-poor soils, but lose this advantage with increasing nitrogen supply. By contrast, the addition of phosphorus, potassium, and other nutrients may benefit such species in low-nutrient environments by enhancing their nitrogen-fixing capacity. We present a global-scale experiment confirming these predictions for nitrogen-fixing legumes (Fabaceae) across 45 grasslands on six continents. Nitrogen addition reduced legume cover, richness, and biomass, particularly in nitrogen-poor soils, while cover of non-nitrogen-fixing plants increased. The addition of phosphorous, potassium, and other nutrients enhanced legume abundance, but did not mitigate the negative effects of nitrogen addition. Increasing nitrogen supply thus has the potential to decrease the diversity and abundance of grassland legumes worldwide regardless of the availability of other nutrients, with consequences for biodiversity, food webs, ecosystem resilience, and genetic improvement of protein-rich agricultural plant species.

Impacts of elevated atmospheric CO2 on arbuscular mycorrhizal fungi and their role in moderating plant allometric partitioning.

Elevated atmospheric CO2 concentration (eCO2) effects on plants depend on several factors including plant photosynthetic physiology (e.g. C3, C4), soil nutrient availability and plants' co-evolved soil-dwelling fungal symbionts, namely arbuscular mycorrhizal (AM) fungi. Complicated interactions among these components will determine the outcomes for plants. Therefore, clearer understanding is needed of how plant growth and nutrient uptake, along with root-colonising AM fungal communities, are simultaneously impacted by eCO2. We conducted a factorial growth chamber experiment with a C3 and a C4 grass species (± AM fungi and ± eCO2). We found that eCO2 increased plant biomass allocation towards the roots, but only in plants without AM fungi, potentially associated with an eCO2-driven increase in plant nutrient requirements. Furthermore, our data suggest a difference in the identities of root-colonising fungal taxa between ambient CO2 and eCO2 treatments, particularly in the C4 grass species, although this was not statistically significant. As AM fungi are ubiquitous partners of grasses, their response to increasing atmospheric CO2 is likely to have important consequences for how grassland ecosystems respond to global change.

Opposing community assembly patterns for dominant and nondominant plant species in herbaceous ecosystems globally.

Biotic and abiotic factors interact with dominant plants-the locally most frequent or with the largest coverage-and nondominant plants differently, partially because dominant plants modify the environment where nondominant plants grow. For instance, if dominant plants compete strongly, they will deplete most resources, forcing nondominant plants into a narrower niche space. Conversely, if dominant plants are constrained by the environment, they might not exhaust available resources but instead may ameliorate environmental stressors that usually limit nondominants. Hence, the nature of interactions among nondominant species could be modified by dominant species. Furthermore, these differences could translate into a disparity in the phylogenetic relatedness among dominants compared to the relatedness among nondominants. By estimating phylogenetic dispersion in 78 grasslands across five continents, we found that dominant species were clustered (e.g., co-dominant grasses), suggesting dominant species are likely organized by environmental filtering, and that nondominant species were either randomly assembled or overdispersed. Traits showed similar trends for those sites (<50%) with sufficient trait data. Furthermore, several lineages scattered in the phylogeny had more nondominant species than expected at random, suggesting that traits common in nondominants are phylogenetically conserved and have evolved multiple times. We also explored environmental drivers of the dominant/nondominant disparity. We found different assembly patterns for dominants and nondominants, consistent with asymmetries in assembly mechanisms. Among the different postulated mechanisms, our results suggest two complementary hypotheses seldom explored: (1) Nondominant species include lineages adapted to thrive in the environment generated by dominant species. (2) Even when dominant species reduce resources to nondominant ones, dominant species could have a stronger positive effect on some nondominants by ameliorating environmental stressors affecting them, than by depleting resources and increasing the environmental stress to those nondominants. These results show that the dominant/nondominant asymmetry has ecological and evolutionary consequences fundamental to understand plant communities.

Age differences in item selection behaviors and subsequent memory for new foreign language vocabulary: Evidence for a region of proximal learning heuristic.

We examined younger and older adults' item selection behaviors to assess heuristics for self-regulating learning of English meanings of Chinese characters varying widely in figural complexity. Two study-test trials were used to assess whether (a) item selection behaviors on the first study opportunity would show evidence for a difficulty-based heuristic as posited by Metcalfe's (2002) region of proximal learning (RPL) theory, or alternatively, influences of habitual English-language reading order (i.e. left-to-right, top-to-bottom); (b) whether second-trial selection behaviors were better predicted by RPL or by the discrepancy reduction model (DRM; Dunlosky & Hertzog, 1998); and (c) whether Trial 1 test performance would alter Trial 2 study in a manner predicted by RPL. DRM stipulates people select any item for study at Trial 2 that was not previously recalled. RPL states that people study only items they believe they can learn, avoiding complex items above their subjective RPL. Stimuli were 36 Chinese-English vocabulary pairs randomly presented in six 2 × 3-element grids. Both habitual reading order and stimulus complexity at Trial 1 affected order of study, with participants of both age groups manifesting a preference to study less complex characters. However, older adults showed larger effects of stimulus complexity whereas younger adults had larger effects of habitual reading order. At Trial 2, older adults showed a greater tendency to avoid studying the most complex Chinese characters, consistent with RPL, which contributed to their lower rates of vocabulary acquisition. Older adults' more conservative RPLs appeared to constrain their degree of self-regulated learning. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

General destabilizing effects of eutrophication on grassland productivity at multiple spatial scales.

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities.

Microbial processing of plant remains is co-limited by multiple nutrients in global grasslands.

Microbial processing of aggregate-unprotected organic matter inputs is key for soil fertility, long-term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro- and micro-nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak-season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains.

Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality.

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands-those with both species-rich local communities (α-diversity) and large compositional differences among localities (ß-diversity)-had higher levels of multifunctionality. Moreover, α- and ß-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities.

Asynchrony among local communities stabilises ecosystem function of metacommunities.

Temporal stability of ecosystem functioning increases the predictability and reliability of ecosystem services, and understanding the drivers of stability across spatial scales is important for land management and policy decisions. We used species-level abundance data from 62 plant communities across five continents to assess mechanisms of temporal stability across spatial scales. We assessed how asynchrony (i.e. different units responding dissimilarly through time) of species and local communities stabilised metacommunity ecosystem function. Asynchrony of species increased stability of local communities, and asynchrony among local communities enhanced metacommunity stability by a wide range of magnitudes (1-315%); this range was positively correlated with the size of the metacommunity. Additionally, asynchronous responses among local communities were linked with species' populations fluctuating asynchronously across space, perhaps stemming from physical and/or competitive differences among local communities. Accordingly, we suggest spatial heterogeneity should be a major focus for maintaining the stability of ecosystem services at larger spatial scales.

The Impact of Presentation Format on Younger and Older Adults' Self-Regulated Learning.

Background/Study Context: Self-regulated learning involves deciding what to study and for how long. Debate surrounds whether individuals' selections are influenced more by item complexity, point values, or if instead people select in a left-to-right reading order, ignoring item complexity and value. The present study manipulated whether point values and presentation format favored selection of simple or complex Chinese-English pairs to assess the impact on younger and older adults' selection behaviors. METHODS: One hundred and five younger (Mage = 20.26, SD = 2.38) and 102 older adults (Mage = 70.28, SD = 6.37) participated in the experiment. Participants studied four different 3 × 3 grids (two per trial), each containing three simple, three medium, and three complex Chinese-English vocabulary pairs presented in either a simple-first or complex-first order, depending on condition. Point values were assigned in either a 2-4-8 or 8-4-2 order so that either simple or complex items were favored. RESULTS: Points did not influence the order in which either age group selected items, whereas presentation format did. Younger and older adults selected more simple or complex items when they appeared in the first column. However, older adults selected and allocated more time to simpler items but recalled less overall than did younger adults. Memory beliefs and working memory capacity predicted study time allocation, but not item selection, behaviors. CONCLUSION: Presentation format must be considered when evaluating which theory of self-regulated learning best accounts for younger and older adults' study behaviors and whether there are age-related differences in self-regulated learning. The results of the present study combine with others to support the importance of also considering the role of external factors (e.g., working memory capacity and memory beliefs) in each age group's self-regulated learning decisions.

The role of font size and font style in younger and older adults' predicted and actual recall performance.

We examined how font sizes (18pt., 48 pt.) and font styles (regular, italic, bold) influenced younger and older adults' judgments of learning (JOLs) and recall. In Experiment 1 younger adults gave higher JOLs and obtained higher recall than older adults. However, JOLs and recall varied for both age groups as a function of font size and font style manipulations despite a tendency for both groups to predict higher recall for items in large and in regular and italic styles than for small and bold fonts and achieve higher recall for regular than italic or bold items. No age differences were found in relative accuracy, with near-perfect calibration in absolute accuracy for younger and older adults. Experiment 2 presented a description of Experiment 1 and asked participants to predict recall for the various font size/style combinations. Younger and older adults predicted higher recall for large than small font items, regardless of font style, and higher recall for bold than regular or italic styles, regardless of font size. Memory predictions did not align across experiments, suggesting that memory beliefs combine with processing fluency to affect JOLs and recall.

Age-related associative memory deficits in value-based remembering: The contribution of agenda-based regulation and strategy use.

Value-based remembering in free-recall tasks may be spared from the typical age-related cognitive decline observed for episodic memory. However, it is unclear whether value-based remembering for associative information is also spared from age-related cognitive decline. The current experiments evaluated the contribution of agenda-based based regulation and strategy use during study to age differences and similarities in value-based remembering of associative information. Participants studied word pairs (Experiments 1-2) or single words (Experiment 2) slated with different point values by moving a mouse controlled cursor to different spatial locations to reveal either items for study or the point value associated with remembering each item. Some participants also provided strategy reports for each item. Younger and older adults allocated greater time to studying high- than low-valued information, reported using normatively effective encoding strategies to learn high-valued pairs, and avoided study of low-valued pairs. As a consequence, both age groups selectively remembered more high- than low-valued items. Despite nearly identical regulatory behavior, an associative memory deficit for older adults was present for high-valued pairs. Age differences in value-based remembering did not occur when the materials were word lists. Fluid intelligence also moderated the effectiveness of older adults' strategy use for high-valued pairs (Experiment 2). These results suggest that age differences in associative value-based remembering may be due to some older adults' gleaning less benefit from using normatively effective encoding strategies rather than age differences in metacognitive self-regulation per se.

The role of warnings in younger and older adults' retrieval-induced forgetting.

Retrieval-induced forgetting (RIF) is a phenomenon wherein practicing recalling some items impairs recall of semantically related, unpracticed items. Two experiments examined whether explicitly warning older (Experiment 1) and younger adults (Experiments 1 and 2) about RIF at different times during two exposures to the retrieval-practice paradigm would affect participants' forgetting. Participants in both experiments were either warned before encoding, retrieval-practice, recall, or not at all. The warning was combined with integration instructions in Experiment 2. Warnings did not reduce forgetting in either age group in Experiment 1. Forgetting increased across exposures in most cases and older adults experienced more forgetting than did younger adults. Combining integration instructions with the warning also did not reduce younger adults' forgetting relative to baseline conditions in Experiment 2. Results indicate that both younger and older adults are susceptible to retrieval-induced forgetting and that raising awareness of the phenomenon increases rather than decreases forgetting rates.

Seasonality and facilitation drive tree establishment in a semi-arid floodplain savanna.

A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree-grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C4 grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C4 grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C4 grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.

The role of feedback and dot presentation format in younger and older adults' number estimation.

Numerosity estimation, the rapid assessment of the number of items in a visual scene, is historically inaccurate. We assessed whether providing feedback regarding the correct numerosity on either 0%, 50%, or 100% of the trials would affect younger and older adults' estimation accuracy for randomized, clustered (i.e., groups of 3 or 7 dots), and stacked (i.e., column) dot formats. Participants provided estimates and confidence ratings in six blocks, each containing 48 trials (16 numerosities shown in each format). Feedback frequency was manipulated between participants during blocks 1-4; no feedback was provided during blocks 5 and 6, which contained old and new numerosities and previously estimated presentations rotated 90°. Estimation accuracy was age equivalent across blocks despite younger adults initially being more accurate than older adults. Feedback improved both age groups' accuracy. Stacked presentations were most accurately estimated but were more likely to be over-estimated than clustered and randomized presentations. Older adults gave lower confidence ratings than younger adults despite both age groups showing increased confidence across blocks, for more structured presentation formats, and as feedback frequency increased. These results expand our understanding of the role of presentation format and feedback in producing age equivalence or age-related differences in numerosity estimation.