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The radiation of a lineage and its rise to ecological dominance are distinct phenomena and driven by different processes. For example, paleoecological data has been used to show that the Cretaceous angiosperm radiation did not coincide with their rise to dominance. Using a phylogenetic approach, we here explored the evolution of C4 grasses and evaluated whether the diversification of this group and its rise to ecological dominance in the late Miocene were decoupled. We assembled a matrix including 675 grass species of the PACMAD clade and 2784 characters (ITS and ndhF ) to run a molecular dating analysis using three fossils as reference calibrations. We coded species as C sub( 3) vs. C sub( 4) and reconstructed ancestral states under maximum likelihood. We used the program BiSSE to test whether rates of diversification are correlated with photosynthetic pathway and whether the radiation of C sub( 4) lineages preceded or coincided with their rise to ecological dominance from ~10 Ma. C sub( 4) grass lineages first originated around 35 Ma at the time of the Eocene-Oligocene transition. Accelerated diversification of C4 lineages did not coincide with their rise to ecological dominance. C sub( 4)-dominated grasslands have expanded only since the Late Miocene and Pliocene. The initial diversification of their biotic elements can be tracked back as far as the Eocene-Oligocene transition. We suggest that shifts in taxonomic diversification and ecological dominance were stimulated by different factors, as in the case of the early angiosperms in the Cretaceous.

Analysis of multi-year nutrient enrichment experiments carried out on 45 global grassland sites show that an addition of an increasing number of nutrients leads to a reduction in plant species diversity, and competition for multiple belowground resources promotes plant species diversity. The roots of species coexistence Theory suggests that the presence of multiple limiting resources within an ecosystem allows for trade-offs between species, promoting the potential for coexistence. William Harpole and colleagues test this theory in plant communities, using data from the international Nutrient Network collaboration. They compile data from multi-year nutrient enrichment experiments carried out on 45 grassland sites on five continents to show that the addition of an increasing number of nutrients leads to a reduction in plant species diversity. The findings suggest that competition for below-ground resources promotes plant species diversity. Niche dimensionality provides a general theoretical explanation for biodiversity—more niches, defined by more limiting factors, allow for more ways that species can coexist 1 . Because plant species compete for the same set of limiting resources, theory predicts that addition of a limiting resource eliminates potential trade-offs, reducing the number of species that can coexist 2 . Multiple nutrient limitation of plant production is common and therefore fertilization may reduce diversity by reducing the number or dimensionality of belowground limiting factors. At the same time, nutrient addition, by increasing biomass, should ultimately shift competition from belowground nutrients towards a one-dimensional competitive trade-off for light 3 . Here we show that plant species diversity decreased when a greater number of limiting nutrients were added across 45 grassland sites from a multi-continent experimental network 4 . The number of added nutrients predicted diversity loss, even after controlling for effects of plant biomass, and even where biomass production was not nutrient-limited. We found that elevated resource supply reduced niche dimensionality and diversity and increased both productivity 5 and compositional turnover. Our results point to the importance of understanding dimensionality in ecological systems that are undergoing diversity loss in response to multiple global change factors.

Grazing exclusion may lead to biodiversity loss and homogenization of naturally heterogeneous and species-rich grassland ecosystems, and these effects may cascade to higher trophic levels and ecosystem properties. Although grazing exclusion has been studied elsewhere, the consequences of alleviating the disturbance regime in grassland ecosystems remain unclear. In this paper, we present results of the first five years of an experiment in native grasslands of southern Brazil. Using a randomized block experimental design, we examined the effects of three grazing treatments on plant and arthropod communities: (i) deferred grazing (i.e., intermittent grazing), (ii) grazing exclusion and (iii) a control under traditional continuous grazing, which were applied to 70 x 70 m experimental plots, in six regionally distributed blocks. We evaluated plant community responses regarding taxonomic and functional diversity (life-forms) in separate spatial components: alpha (1 x 1 m subplots), beta, and gamma (70 x 70 m plots), as well as the cascading effects on arthropod high-taxa. By estimating effect sizes (treatments vs. control) by bootstrap resampling, both deferred grazing and grazing exclusion mostly increased vegetation height, plant biomass and standing dead biomass. The effect of grazing exclusion on plant taxonomic diversity was negative. Conversely, deferred grazing increased plant taxonomic diversity, but both treatments reduced plant functional diversity. Reduced grazing pressure in both treatments promoted the break of dominance by prostrate species, followed by fast homogenization of vegetation structure towards dominance of ligneous and erect species. These changes in the plant community led to increases in high-taxa richness and abundance of vegetation-dwelling arthropod groups under both treatments, but had no detectable effects on epigeic arthropods. Our results indicate that decision-making regarding the conservation of southern Brazil grasslands should include both intensive and alleviated levels of grazing management, but not complete grazing exclusion, to maximize conservation results when considering plant and arthropod communities.

Sorghum is emerging as an excellent genetic model for the design of C₄ grass bioenergy crops. Annual energy Sorghum hybrids also serve as a source of biomass for bioenergy production. Elucidation of Sorghum’s flowering time gene regulatory network, and identification of complementary alleles for photoperiod sensitivity, enabled large-scale generation of energy Sorghum hybrids for testing and commercial use. Energy Sorghum hybrids with long vegetative growth phases were found to accumulate more than twice as much biomass as grain Sorghum, owing to extended growing seasons, greater light interception, and higher radiation use efficiency. High biomass yield, efficient nitrogen recycling, and preferential accumulation of stem biomass with low nitrogen content contributed to energy Sorghum’s elevated nitrogen use efficiency. Sorghum’s integrated genetics-genomics-breeding platform, diverse germplasm, and the opportunity for annual testing of new genetic designs in controlled environments and in multiple field locations is aiding fundamental discovery, and accelerating the improvement of biomass yield and optimization of composition for biofuels production. Recent advances in wide hybridization between Sorghum and other C₄ grasses could allow the deployment of improved genetic designs of annual energy Sorghums in the form of wide-hybrid perennial crops. The current trajectory of energy Sorghum genetic improvement indicates that it will be possible to sustainably produce biofuels from C₄ grass bioenergy crops that are cost competitive with petroleum-based transportation fuels.

Genomic DNA base composition (GC content) is predicted to significantly affect genome functioning and species ecology. Although several hypotheses have been put forward to address the biological impact of GC content variation in microbial and vertebrate organisms, the biological significance of GC content diversity in plants remains unclear because of a lack of sufficiently robust genomic data. Using flow cytometry, we report genomic GC contents for 239 species representing 70 of 78 monocot families and compare them with genomic characters, a suite of life history traits and climatic niche data using phylogeny-based statistics. GC content of monocots varied between 33.6% and 48.9%, with several groups exceeding the GC content known for any other vascular plant group, highlighting their unusual genome architecture and organization. GC content showed a quadratic relationship with genome size, with the decreases in GC content in larger genomes possibly being a consequence of the higher biochemical costs of GC base synthesis. Dramatic decreases in GC content were observed in species with holocentric chromosomes, whereas increased GC content was documented in species able to grow in seasonally cold and/or dry climates, possibly indicating an advantage of GC-rich DNA during cell freezing and desiccation. We also show that genomic adaptations associated with changing GC content might have played a significant role in the evolution of the Earth’s contemporary biota, such as the rise of grass-dominated biomes during the mid-Tertiary. One of the major selective advantages of GC-rich DNA is hypothesized to be facilitating more complex gene regulation. Significance Our large-scale survey of genomic nucleotide composition across monocots has enabled the first rigorous testing, to our knowledge, of its biological significance in plants. We show that genomic DNA base composition (GC content) is significantly associated with genome size and holocentric chromosomal structure. GC content may also have deep ecological relevance, because changes in GC content may have played a significant role in the evolution of Earth’s biota, especially the rise of grass-dominated biomes during the mid-Tertiary. The discovery of several groups with very unusual GC contents highlights the need for in-depth analysis to uncover the full extent of genomic diversity. Furthermore, our stratified sampling method of distribution data and quantile regression-like logic of phylogenetic analyses may find wider applications in the analysis of spatially heterogeneous data.

The evolution of grasses using C₄ photosynthesis and their sudden rise to ecological dominance 3 to 8 million years ago is among the most dramatic examples of biome assembly in the geological record. A growing body of work suggests that the patterns and drivers of C₄ grassland expansion were considerably more complex than originally assumed. Previous research has benefited substantially from dialog between geologists and ecologists, but current research must now integrate fully with phylogenetics. A synthesis of grass evolutionary biology with grassland ecosystem science will further our knowledge of the evolution of traits that promote dominance in grassland systems and will provide a new context in which to evaluate the relative importance of C₄ photosynthesis in transforming ecosystems across large regions of Earth.

C ₄ photosynthesis is a series of anatomical and biochemical modifications to the typical C ₃ pathway that increases the productivity of plants in warm, sunny, and dry conditions. Despite its complexity, it evolved more than 62 times independently in flowering plants. However, C ₄ origins are absent from most plant lineages and clustered in others, suggesting that some characteristics increase C ₄ evolvability in certain phylogenetic groups. The C ₄ trait has evolved 22–24 times in grasses, and all origins occurred within the PACMAD clade, whereas the similarly sized BEP clade contains only C ₃ taxa. Here, multiple foliar anatomy traits of 157 species from both BEP and PACMAD clades are quantified and analyzed in a phylogenetic framework. Statistical modeling indicates that C ₄ evolvability strongly increases when the proportion of vascular bundle sheath (BS) tissue is higher than 15%, which results from a combination of short distance between BS and large BS cells. A reduction in the distance between BS occurred before the split of the BEP and PACMAD clades, but a decrease in BS cell size later occurred in BEP taxa. Therefore, when environmental changes promoted C ₄ evolution, suitable anatomy was present only in members of the PACMAD clade, explaining the clustering of C ₄ origins in this lineage. These results show that key alterations of foliar anatomy occurring in a C ₃ context and preceding the emergence of the C ₄ syndrome by millions of years facilitated the repeated evolution of one of the most successful physiological innovations in angiosperm history.

Background Puccinellia tenuiflora is the most saline-alkali tolerant plant in the Songnen Plain, one of the three largest soda saline-alkali lands worldwide. Here, we investigated the physicochemical properties of saline-alkali soils from the Songnen Plain and sequenced the transcriptomes of germinated P. tenuiflora seedlings under long-term treatment (from seed soaking) with saline-alkali soil extracts. Results We found that the soils from Songnen Plain were reasonably rich in salts and alkali; moreover, the soils were severely deficient in nitrogen [N], phosphorus [P], potassium [K] and several other mineral elements. This finding demonstrated that P. tenuiflora can survive from not only high saline-alkali stress but also a lack of essential mineral elements. To explore the saline-alkali tolerance mechanism, transcriptional analyses of P. tenuiflora plants treated with water extracts from the saline-alkali soils was performed. Interestingly, unigenes involved in the uptake of N, P, K and the micronutrients were found to be significantly upregulated, which indicated the existence of an efficient nutrition-uptake system in P. tenuiflora . Compared with P. tenuiflora , the rice Oryza sativa was hypersensitive to saline-alkali stress. The results obtained using a noninvasive microtest techniques confirmed that the uptake of NO 3 - and NH 4 + and the regulatory flux of Na + and H + were significantly higher in the roots of P. tenuiflora than in those of O. sativa . In the corresponding physiological experiments, the application of additional nutrition elements significantly eliminated the sensitive symptoms of rice to saline-alkali soil extracts. Conclusions Our results imply that the survival of P. tenuiflora in saline-alkali soils is due to a combination of at least two regulatory mechanisms and the high nutrient uptake capacity of P. tenuiflora plays a pivotal role in its adaptation to those stress. Taken together, our results highlight the role of nutrition uptake in saline-alkali stress tolerance in plants.

Warming, watering and elevated atmospheric CO₂-concentration effects have been extensively studied separately; however, their combined impact on plants is not well understood. In the current research, we examined plant growth and physiological responses of three dominant species from the Eurasian Steppe with different functional traits to a combination of elevated CO₂, high temperature, and four simulated precipitation patterns. Elevated CO₂ stimulated plant growth by 10.8–41.7 % for a C₃ leguminous shrub, Caragana microphylla, and by 33.2–52.3 % for a C₃ grass, Stipa grandis, across all temperature and watering treatments. Elevated CO₂, however, did not affect plant biomass of a C₄ grass, Cleistogenes squarrosa, under normal or increased precipitation, whereas a 20.0–69.7 % stimulation of growth occurred with elevated CO₂ under drought conditions. Plant growth was enhanced in the C₃ shrub and the C₄ grass by warming under normal precipitation, but declined drastically with severe drought. The effects of elevated CO₂ on leaf traits, biomass allocation and photosynthetic potential were remarkably species-dependent. Suppression of photosynthetic activity, and enhancement of cell peroxidation by a combination of warming and severe drought, were partly alleviated by elevated CO₂. The relationships between plant functional traits and physiological activities and their responses to climate change were discussed. The present results suggested that the response to CO₂ enrichment may strongly depend on the response of specific species under varying patterns of precipitation, with or without warming, highlighting that individual species and multifactor dependencies must be considered in a projection of terrestrial ecosystem response to climatic change.

Grasslands cover more than 20% of the Earth's terrestrial surface, and their rise to dominance is one of the most dramatic events of biome evolution in Earth history. Grasses possess two main photosynthetic pathways: the C₃ pathway that is typical of most plants and a specialized C₄ pathway that minimizes photorespiration and thus increases photosynthetic performance in high-temperature and/or low-CO₂ environments. C₄ grasses dominate tropical and subtropical grasslands and savannas, and C₃ grasses dominate the world's cooler temperate grassland regions. This striking pattern has been attributed to C₄ physiology, with the implication that the evolution of the pathway enabled C₄ grasses to persist in warmer climates than their C₃ relatives. We combined geospatial and molecular sequence data from two public archives to produce a 1,230-taxon phylogeny of the grasses with accompanying climate data for all species, extracted from more than 1.1 million herbarium specimens. Here we show that grasses are ancestrally a warm-adapted clade and that C₄ evolution was not correlated with shifts between temperate and tropical biomes. Instead, 18 of 20 inferred C₄ origins were correlated with marked reductions in mean annual precipitation. These changes are consistent with a shift out of tropical forest environments and into tropical woodland/savanna systems. We conclude that C₄ evolution in grasses coincided largely with migration out of the understory and into open-canopy environments. Furthermore, we argue that the evolution of cold tolerance in certain C₃ lineages is an overlooked innovation that has profoundly influenced the patterning of grassland communities across the globe.