Large and non-spherical seeds are less likely to form a persistent soil seed bank.

There is some evidence that seed traits can affect the long-term persistence of seeds in the soil. However, findings on this topic have differed between systems. Here, we brought together a worldwide database of seed persistence data for 1474 species to test the generality of seed mass-shape-persistence relationships. We found a significant trend for low seed persistence to be associated with larger and less spherical seeds. However, the relationship varied across different clades, growth forms and species ecological preferences. Specifically, relationships of seed mass-shape-persistence were more pronounced in Poales than in other order clades. Herbaceous species that tend to be found in sites with low soil sand content and precipitation have stronger relationships between seed shape and persistence than in sites with higher soil sand content and precipitation. For the woody plants, the relationship between persistence and seed morphology was stronger in sites with high soil sand content and low precipitation than in sites with low soil sand content and higher precipitation. Improving the ability to predict the soil seed bank formation process, including burial and persistence, could benefit the utilization of seed morphology-persistence relationships in management strategies for vegetation restoration and controlling species invasion across diverse vegetation types and environments.

Effect of grazing and nitrogen addition on the occurrence of species with different seed masses in alpine meadows on the Tibet Plateau.

Seed mass (SM) is a core functional trait of plant species. Thus, information of the effect of grazing and nitrogen addition on the occurrence of species with different SMs can help us understand the influence of grazing and fertilization on survival of species and community assembly. In alpine meadows with different grazing and nitrogen addition treatments on the eastern Tibet Plateau, we measured SM and plant height (H) of the most common plants and conducted a series of quadrat surveys. Overall, grazing promoted survival of small-seeded species, while fertilization suppressed their survival. At the community level, moderate grazing reduced the average community weighted mean (CWM) of SM and increased the coefficient of variation (CV) of SM. However, there was no significant difference between the functional diversity (FDrao, calculated as Rao's index) of SM in grazed and nongrazed meadows. Nitrogen addition significantly increased the CWM of SM, had a marginally significant effect on FDrao of SM, but had no significant effect on CV of SM. The impact of gazing and fertilization on occurrence of species with different SMs can be explained by their effect on vegetation height. From the perspective of SM selection, our study helps clarify the mechanism of species diversity loss due to fertilization, and that of species diversity increase by moderate grazing.

Regional gradients in intraspecific seed mass variation are associated with species biotic attributes and niche breadth.

Quantifying intraspecific trait variation (ITV) is crucial for understanding species local adaptation and regional distribution. Intraspecific seed mass variation (ITVsm) is expected to vary with environmental gradients or co-vary with related biotic attributes, but these relationships are not well known in a multispecies space. We performed interspecific and phylogenetic analyses to evaluate the relative power of three species biotic attributes and four niche breadth traits in explaining ITVsm variation for 434 eastern Qinghai-Tibetan species. We showed a positive relationship between species' ITVsm and their niche breadth in the light, moisture and disturbance dimensions, supporting the idea that high ITV allows species to match their traits to different habitat conditions and thus to distribute in a wide range of environments. However, we did find significant direct effect of species' thermal niche on individual seed mass variation. Meanwhile, we showed significant effects of seed dispersal mode, but not of life form and pollination type, on ITVsm. This suggests that the covariation or co-evolution between seed and disperser was related to the pattern and magnitude of ITVsm, but not to plant lifespan, the quality and allocation pattern of available resources and the availability of pollination vector. Lastly, all multivariate models showed a significant combined contribution of species' biotic attributes and niche breadth to their ITVsm, implying that intrinsic biotic limitations and extrinsic abiotic pressures may operate simultaneously in controlling regional-scale intraspecific seed development.

Geographic Variation in the Petiole-Lamina Relationship of 325 Eastern Qinghai-Tibetan Woody Species: Analysis in Three Dimensions.

The petiole-lamina relationship is central to the functional tradeoff between photosynthetic efficiency and the support/protection cost. Understanding environmental gradients in the relationship and its underlying mechanisms remains a critical challenge for ecologists. We investigated the possible scaling of the petiole-lamina relationships in three dimensions, i.e., petiole length (PL) vs. lamina length (LL), petiole cross sectional area (PCA) vs. lamina area (LA), and petiole mass (PM) vs. lamina mass (LM), for 325 Qinghai-Tibetan woody species, and examined their relation to leaf form, altitude, climate, and vegetation types. Both crossspecies analysis and meta-analysis showed significantly isometric, negatively allometric, and positively allometric scaling of the petiole-lamina relationships in the length, area, and mass dimensions, respectively, reflecting an equal, slower, and faster variation in the petiole than in the lamina in these trait dimensions. Along altitudinal gradients, the effect size of the petiole-lamina relationship decreased in the length and mass dimensions but increased in the area dimension, suggesting the importance of enhancing leaf light-interception and nutrient transport efficiency in the warm zones in petiole development, but enhancing leaf support/protection in the cold zones. The significant additional influences of LA, LM, and LA were observed on the PL-LL, PCA-LA, and PM-LM relationships, respectively, implying that the single-dimension petiole trait is affected simultaneously by multidimensional lamina traits. Relative to simple-leaved species, the presence of petiolule in compound-leaved species can increase both leaf light interception and static gravity loads or dynamic drag forces on the petiole, leading to lower dependence of PL variation on LL variation, but higher biomass allocation to the petiole. Our study highlights the need for multidimension analyses of the petiole-lamina relationships and illustrates the importance of plant functional tradeoffs and the change in the tradeoffs along environmental gradients in determining the relationships.

Effect of Nitrogen Addition on Selection of Germination Trait in an Alpine Meadow on the Tibet Plateau.

Seed germination requirements may determine the kinds of habitat in which plants can survive. We tested the hypothesis that nitrogen (N) addition can change seed germination trait-environmental filter interactions and ultimately redistribute seed germination traits in alpine meadows. We determined the role of N addition on germination trait selection in an alpine meadow after N addition by combining a 3-year N addition experiment in an alpine meadow and laboratory germination experiments. At the species level, germination percentage, germination rate (speed) and breadth of temperature niche for germination (BTN) were positively related to survival of a species in the fertilized community. In addition, community-weighted means of germination percentage, germination rate, germination response to alternating temperature and BTN increased. However, germination response to wet-cold storage (cold stratification) and functional richness of germination traits was lower in alpine meadows with high-nitrogen addition than in those with no, low and medium N addition. Thus, N addition had a significant influence on environmental filter-germination trait interactions and generated a different set of germination traits in the alpine meadow. Further, the effect of N addition on germination trait selection by environmental filters was amount-dependent. Low and medium levels of N addition had less effect on redistribution of germination traits than the high level.

Disrupting Reconsolidation by Systemic Inhibition of mTOR Kinase via Rapamycin Reduces Cocaine-Seeking Behavior.

Drug addiction is considered maladaptive learning, and drug-related memories aroused by the presence of drug related stimuli (drug context or drug-associated cues) promote recurring craving and reinstatement of drug seeking. The mammalian target of rapamycin signaling pathway is involved in reconsolidation of drug memories in conditioned place preference and alcohol self-administration (SA) paradigms. Here, we explored the effect of mTOR inhibition on reconsolidation of addiction memory using cocaine self-administration paradigm. Rats received intravenous cocaine self-administration training for 10 consecutive days, during which a light/tone conditioned stimulus was paired with each cocaine infusion. After acquisition of the stable cocaine self-administration behaviors, rats were subjected to nosepoke extinction (11 days) to extinguish their behaviors, and then received a 15 min retrieval trial with or without the cocaine-paired tone/light cue delivery or without. Immediately or 6 h after the retrieval trial, rapamycin (10 mg/kg) was administered intraperitoneally. Finally, cue-induced reinstatement, cocaine-priming-induced reinstatement and spontaneous recovery of cocaine-seeking behaviors were assessed in rapamycin previously treated animals, respectively. We found that rapamycin treatment immediately after a retrieval trial decreased subsequent reinstatement of cocaine seeking induced by cues or cocaine itself, and these effects lasted at least for 28 days. In contrast, delayed intraperitoneal injection of rapamycin 6 h after retrieval or rapamycin injection without retrieval had no effects on cocaine-seeking behaviors. These findings indicated that mTOR inhibition within the reconsolidation time-window impairs the reconsolidation of cocaine associated memory, reduces cocaine-seeking behavior and prevents relapse, and these effects are retrieval-dependent and temporal-specific.

The seed germination spectrum of alpine plants: a global meta-analysis.

Assumptions about the germination ecology of alpine plants are presently based on individual species and local studies. A current challenge is to synthesise, at the global level, the alpine seed ecological spectrum. We performed a meta-analysis of primary data from laboratory experiments conducted across four continents (excluding the tropics) and 661 species, to estimate the influence of six environmental cues on germination proportion, mean germination time and germination synchrony; accounting for seed morphology (mass, embryo : seed ratio) and phylogeny. Most alpine plants show physiological seed dormancy, a strong need for cold stratification, warm-cued germination and positive germination responses to light and alternating temperatures. Species restricted to the alpine belt have a higher preference for warm temperatures and a stronger response to cold stratification than species whose distribution extends also below the treeline. Seed mass, embryo size and phylogeny have strong constraining effects on germination responses to the environment. Globally, overwintering and warm temperatures are key drivers of germination in alpine habitats. The interplay between germination physiology and seed morphological traits further reflects pressures to avoid frost or drought stress. Our results indicate the convergence, at the global level, of the seed germination patterns of alpine species.

Allocation Strategies for Seed Nitrogen and Phosphorus in an Alpine Meadow Along an Altitudinal Gradient on the Tibetan Plateau.

Nitrogen (N) and phosphorus (P) play important roles in many aspects of plant biology. The allocation of N and P in plant vegetative organs (i.e., leaves, stems, and fine roots) is critical to the regulation of plant growth and development. However, how these elements are allocated in seeds is unclear. The aim of this study was to explore the N and P allocation strategies of seeds in an alpine meadow along an altitudinal gradient. We measured the seed N and P contents of 253 herbaceous species in 37 families along an altitudinal gradient (2,000-4,200 m) in the east Tibetan alpine meadow. The geometric means of seed N and P concentrations and N:P ratios were 34.81 mg g-1, 5.06 mg g-1, and 6.88, respectively. Seed N and P concentrations varied across major taxonomic groups and among different altitude zones. N:P ratios showed no significant variations among different taxonomic groups with the exception of N-fixing species. The numerical value of the scaling exponent of seed N vs. P was 0.73, thus approaching 3/4, across the entire data set, but varied significantly across major taxonomic groups. In addition, the numerical value of the scaling exponent of N vs. P declined from 0.88 in the high altitude zone to 0.63 in the low altitude zone. These results indicate that the variations in the numerical value of the scaling exponent governing the seed N vs. P scaling relationship varies as a function of major taxonomic groups and among different altitude zones. We speculate that this variation reflects different adaptive strategies for survival and germination in an alpine meadow. If true, the data presented here advance our understanding of plant seed allocation strategies, and have important implications for modeling early plant growth and development.

Silicon addition improves plant productivity and soil nutrient availability without changing the grass:legume ratio response to N fertilization.

Silicon (Si) plays an important role in plant nutrient capture and absorption, and also promotes plant mechanical strength and light interception in alpine meadows. In this study, we conducted a field experiment to examine the effect of nitrogen (N) application, with (N + Si) and without Si (N-only), on the potential for soil nutrient and the growth of grass and legume plant functional types (PFTs) in an alpine meadow. It was found that N + Si resulted in higher soil nutrient contents, leaf N and P concentrations, abundance and biomass of legume and grass PFTs than N-only. The aboveground biomass of grass (598 g m-2) and legume (12.68 g m-2) PFTs under 600 kg ha-1 ammonium nitrate (NH4NO3) per year addition with Si was significantly higher than that under the same level of N addition without Si (515 and 8.68 g m-2, respectively). The grass:legume biomass ratio did not differ significantly between the N + Si and N-only. This demonstrates that Si enhances N fertilization with apparently little effect on grass:legume ratio and increases plant-available nutrients, indicating that Si is essential for the plant community in alpine meadows.

Influences of nitrogen, phosphorus and silicon addition on plant productivity and species richness in an alpine meadow.

Fertilization, especially with nitrogen (N), increases aboveground primary productivity (APP), but reduces plant species richness at some level. Silicon (Si) fertilization alone, or with addition of N or phosphorus (P), has multiple direct and indirect beneficial effects on plant growth and development, both for individuals and the whole community. This study aimed to examine the effects of Si, N, P, NSi and PSi combinations on APP and species richness of the community and of four functional groups in an alpine meadow. The results showed that plots fertilized with Si in combination with either N or P had higher APP than when fertilized with N or P alone. Addition of N or P increased APP, and the higher APP occurred when the highest level of N was added, indicating co-limitation of N and P, with N being most limiting. Silicon fertilization alone or with addition of N increased the APP of grasses and forbs. Nitrogen addition decreased the community species richness; Si with addition of N alleviated the loss of species richness of the whole community and the forbs group. For the four functional groups, N or P addition increased the species richness of grasses and decreased that of forbs. Our findings highlight the importance of Si in improving APP and alleviating N fertilization-induced biodiversity loss in grasslands, and will help improve our ability to predict community composition and biomass dynamics in alpine meadow ecosystems subject to changing nutrient availability.

Effect of diurnal fluctuating versus constant temperatures on germination of 445 species from the eastern Tibet Plateau.

Germination response to fluctuating temperatures is a mechanism by which seeds detect gaps in vegetation canopies and depth of burial in soil, and it is very important for plants. Thus, studies on the effect of fluctuating temperature on germination at the community level are valuable for understanding community structure and biodiversity maintenance. We determined the effects of two alternating temperatures (5/25°C and 10/20°C) and one constant temperature (15°C) on seed germination of 445 species in a grassland community on the eastern Tibet Plateau. Seed mass was determined for each species, and data on habitat, type of life cycle, altitudinal distribution and functional group (graminoids or forbs) were obtained from the literature. Taking all species into account, alternating temperatures increased germination percentages regardless of amplitude. Overall, species growing in disturbed ground showed a significant germination response to temperature fluctuation, but those living in Alpine/subalpine meadow, forest margin /scrub, marshland and dry sunny slope habitats did not. Species distributed only at high elevations (>2000m) did not show a significant germination response to temperature fluctuation, whereas those occurring at both high and low elevations had a significant positive response. Germination of annuals/biennials was significantly promoted by 5/25°C, but not by 10/20°C, whereas germination of perennials was significantly promoted by both 5/25°C and 10/20°C. Small-seeded species were more likely than large-seeded species to respond positively to fluctuating temperatures. Germination of forbs had a positive response to temperature fluctuation, but germination of graminoids did not. Regeneration ability by seeds for about 36% of the species studied in the grassland can be increased by temperature fluctuation. The differential response among species to alternating vs. constant temperatures helps maintain community structure and biodiversity. A positive germination response to temperature fluctuation can partly explain why there are more forbs in degraded meadows.

Effect of storage conditions on germination of seeds of 489 species from high elevation grasslands of the eastern Tibet Plateau and some implications for climate change.

PREMISE OF THE STUDY: Little is known about the effect of environmental conditions on seed dormancy break at the community level or how it could be affected by climate change. This study tested the effects of storage conditions on germination of 489 species from high-elevation natural grasslands on the Tibet Plateau. METHODS: We stored seeds in dry cold, dry warm, and wet cold environments to test the effect of these conditions on germination. Germination responses were classified with the use of cluster analysis. The effect of phylogeny on germination response to storage conditions was determined. KEY RESULTS: Compared with results of wet cold conditions, storing seeds at dry warm or dry cold conditions decreased the mean community germination percentage by 17.93% and 16.07%, respectively. Storing seeds at dry warm vs. dry cold conditions decreased the community mean germination percentage by 4.61%. The germination response to moisture conditions during storage showed significant phylogenetic patterns, whereas the germination response to storage temperature did not. On the basis of the germinability of seeds of the same species stored under different conditions, germination may increase, decrease, or stay the same. CONCLUSIONS: Within the high-elevation Tibetan grassland community, the different responses in dormancy breaking and germination behavior to the same storage conditions may have implications for understanding how this community might respond to climate change. In particular, some species may increase in the community, whereas others may decrease or remain the same. However, the potential for such change can be detected only by studying the responses of many species.