Haiyan Bu

Seed mass and germination in an alpine meadow on the eastern Tsinghai-Tibet plateau

In this study, we built up a database of 570 species from an alpine meadow on the eastern Tsinghai–Tibet plateau. We examined the correlation of seed mass and germination with phylogeny, habitat and altitude, and the relationship between seed mass and germination. We found that: habitats had no significant effects on seed mass and germinability, which was in accord with the former studies; there was a significant negative correlation between seed mass and altitude, as well as between germinability and altitude, which was opposite to most of the former studies; there was a significant negative correlation between seed mass and germinability, which was in contrast with other studies that have found a positive relationship, and seed mass could explain 24.1% of total variation in germinability; in GLM, family and genus accounted for 43.9% and 83.9% of total variation in seed mass, and 34.1% and 65.4% in germinability, respectively, thus, it was evident that seed mass and germinability were strongly related to phylogeny. We considered that seed mass and germination might be the result of both selective pressures over long-term ecological time and the constraints over long-standing evolutionary history of the taxonomic membership. We suggest that correlates of ecology and phylogeny should be taken into account in comparative studies on seed mass and germination among species.

Community-wide germination strategies in an alpine meadow on the eastern Qinghai-Tibet plateau: phylogenetic and life-history correlates

In this study, we built up a database of 633 species (48 families, 205 genera) from an alpine meadow on the eastern Qinghai-Tibet plateau. Our objective was to assess the effects of phylogenetic and life-history (life form, perenniality, seed size, dispersal strategy and period) background on the community-wide germination strategies. We found that the seeds of shrubs, perennials, and well-dispersed plants, and the smaller seeds germinated more and comparatively earlier. In one-way ANOVAs, phylogenetic groups explained 12% of the variance in GT (mean germination time for all seeds germinated of each species); life-history attributes, such as seed size, dispersal strategy, perenniality and life form explained 10%, 7%, 5%, and 1% respectively, and dispersal period had no significant effect on GT. Multifactorial ANOVAs revealed that the three major factors contributing to differences in GT were phylogenetic relatedness, seed size and dispersal strategy (explained 4%, 5% and 4% of the interspecific variation independently, respectively). Thus, seeds germination strategies were significantly correlated with phylogenetic and life-history relatedness. In addition, phylogenetic relatedness had close associations and interactions with seed size and dispersal strategy. Then, we think phylogeny and life-history attributes could not be considered mutual exclusively. Seed germination, like any other trait, is shaped by the natural history of the species and by the evolutionary history of the lineage. And a large percentage of the variance remained unexplained by our model, which suggested important selective factors or parameters may have been left out of this analysis.

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.

The ecological and evolutionary significance of seed shape and volume for the germination of 383 species on the eastern Qinghai-Tibet plateau

Seed shape and seed volume are functionally important traits which are associated with seed germination. To study the ecological and evolutionary significance of seed shape and volume in relation to germination, a database comprising seed information for 383 species within 50 families from an alpine meadow on the eastern Qinghai-Tibet plateau was compiled. The independent and interactive effects of seed traits on germination and whether these effects are controlled by phylogeny are discussed. The results show the following: (1) Elongated seeds had higher percentage and faster speed of germination than compact seeds whereas smaller seeds tended to have higher germinability compared to larger ones; (2) Seed volume and shape evolved together but had independent effects on seed germination. Seed volume had no significant influence on seed germination among species with compact seeds, but had a significant effect on those among species with elongated seeds; seed germination of larger seeded species was affected by seed shape, but not by seed volume. These results might be related to their predation pressure and ability to germinate from deeper soil layers. Lastly, (3) Phylogenetic conservation plays an important role in the co-evolution of seed traits and germination characteristics. Species with similarly shaped seeds on a large scale might reflect a common selective pressure, but related species within a family might be more dependent on phylogenetic conservation.

Untangling interacting mechanisms of seed mass variation with elevation: insights from the comparison of inter-specific and intra-specific studies on eastern Tibetan angiosperm species

With increasing elevation, seed mass is expected to be either larger for its advantage during seedling establishment in stressful high-elevation environments (“stress-tolerance” mechanism) or smaller due to energy constraints. Based on the combination of inter- and intra-specific analyses on 4,023 populations of 320 Tibetan plant species, we found an overall positive within species but negative among species seed mass–elevation relationship, suggesting that regional seed mass distribution with elevation is affected mainly by the energy-constraint mechanism at among-species level, but by the stress-tolerance mechanism at within-species level. Moreover, both intra- and inter-specific analyses revealed a mass-dependent seed mass variation along elevation gradients: small seeds tended to increase but large seeds tended to decrease with increasing elevation, indicating that the stress-tolerance (respectively energy-constraint) mechanism may exert a stronger effect on elevational distribution of small-seeded (respectively large-seeded) populations or species. The mass-dependent seed mass variation along elevation gradients, however, was absent within woody and zoochorous species, implying substantial available resources and long time for seed development, and the covariation or coevolution between the mass of zoochorous seeds and their dispersers may allow for a (partial) decoupling of species’ seed mass and their intra-specific variation with elevation. Together our results call for more comparative analyses at different taxonomic levels in detecting geographic variation in a trait.

The effects of phylogeny, life-history traits and altitude on the carbon, nitrogen, and phosphorus contents of seeds across 203 species from an alpine meadow

Seed reserves are very important to the earlier growth and survival of plants, and its variation in nutrient contents might make species to form different germination or seedlings growth strategies. Here, 203 species collected from an alpine meadow on the northeastern Qinghai–Tibet plateau were used to test the effects of phylogenetic groups, life-history traits, and altitude on carbon (C), nitrogen (N) and phosphorus (P) contents of seeds across species. The results showed that (1) seeds of Brassicaceae had the highest C content, those of Fabaceae had the highest N content, and Asteraceae had the highest P content, and family explained independently 32.7%, 46.4%, and 17.9% of the variation in C, N, and P contents of seeds, respectively; (2) the smaller seeds tended to have higher C and P contents, and seed mass explained independently 2.5% of variation in C, 4.3% in N, and 8.1% in P contents; (3) N content was explained 1% independently by life form, seeds N content of perennials was significantly higher than that of annals, but seeds C and P contents had non-significant difference between them; (4) seeds of wind-adapted species had higher N and P contents, and dispersal mode explained independently 1.7% of variation in C, 1.6% in N, and 5.6% in P contents; (5) seeds from high altitude had the highest N and P contents, and altitude explained independently 4.3% of the variation in N and 4% in P contents.

The effect of light and seed mass on seed germination of common herbaceous species from the eastern Qinghai‐Tibet Plateau

Smaller seeds might encounter more severe selective pressure than larger ones because they have fewer food reserves and are more easily buried; thus, seed mass can be considered to be directly related to the effect of light on germination. To investigate the effect of light on seed germination and associated seed mass variation within a whole plant community, we compiled germination data for common herbaceous species from an alpine meadow on the eastern Qinghai-Tibet plateau. The results showed the following. (i) Light had a general positive effect on seed germination. Under light, the proportion of species with lower germinability was decreased, mean germination percentage was increased by 20% and the speed of germination was doubled. (ii) Irrespective of light environment, species with medium-sized seeds (seed mass ranging from 0.11 to 0.5 mg) had higher germination percentage and speed when compared with species within the largest seed mass group. (iii) The germination of smaller-seeded species was more dependent on light stimulation than larger-seeded ones. In darkness, the species within the smallest seed mass group had the lowest percentage and speed of germination; however, under light, the species within the largest seed mass group had the lowest percentage and speed of germination. Our results suggested that the germination characteristics and especially seeds’ response to light among species in the alpine meadow might be an adaptation to natural selective pressure.

Seed germinating characteristics of 54 gramineous species in the alpine meadow on the eastern Qinghai-Tibet plateau

The germination experiment was started on March 3, 2004, and seeds were collected from July to October in 2003. We analyzed the percentage of germination, days to first germination, germination period and days to 50% germination. Among the 54 examined species, 26 species exceeded 80% germination, 11 species exceeded 60%–80% germination, 8 exceeded 40%–0%, 5 exceeded 20%–40%, and 4 showed less than 20%. A principal-component analysis revealed that the species were distributed along two statistically independent axes, the first primarily represented the germination rate and the second represented the time of germination process. Based on scores of germination characteristics, cluster analysis of the 54 gramineous species could be divided into 4 distinct groups: rapid, slow, intermediate germinating (germination percentage > 50%), and low germinating (germination percentage < 50%). The meaning of different groups to the vegetation regeneration was discussed.