Global seed dormancy patterns are driven by macroclimate but not fire regime.

Seed dormancy maximizes plant recruitment in habitats with variation in environmental suitability for seedling establishment. Yet, we still lack a comprehensive synthesis of the macroecological drivers of nondormancy and the different classes of seed dormancy: physiological dormancy, morphophysiological dormancy and physical dormancy. We examined current geographic patterns and environmental correlates of global seed dormancy variation. Combining the most updated data set on seed dormancy classes for > 10 000 species with > 4 million georeferenced species occurrences covering all of the world's biomes, we test how this distribution is driven by climate and fire regime. Seed dormancy is prevalent in seasonally cold and dry climates. Physiological dormancy occurs in relatively dry climates with high temperature seasonality (e.g. temperate grasslands). Morphophysiological dormancy is more common in forest-dominated, cold biomes with comparatively high and evenly distributed precipitation. Physical dormancy is associated with dry climates with strong seasonal temperature and precipitation fluctuations (e.g. deserts and savannas). Nondormancy is associated with stable, warm and wetter climates (e.g. tropical rain forest). Pyroclimate had no significant effect on the distribution of seed dormancy. The environmental drivers considered in this study had a comparatively low predictive power, suggesting that macroclimate is just one of several global drivers of seed dormancy.

Utkin et al.'s dataset on specific leaf area.

Specific leaf area (SLA; one-sided leaf area per unit of dry mass) is a key trait indicating plant growth strategy and its responses to changing environments. Despite its high relevance for ecological research and recent efforts to mobilize the trait data, information on SLA in many plant lineages and biogeographic regions is still underrepresented. To assist in closing this gap, we translated and digitized a large dataset on SLA titled the Surface areas of forest plants by Anatoly I. Utikin, Lyudmila S. Ermolova, and Irina A. Utkina, published in Russian in 2008 (Nakua, Moscow, Russia) and previously not accessible to the great majority of people in the international research community. The book contains a compendium of SLA values collected by A. Utkin and his colleagues published between 1918 and 2006 containing research on ~1100 gymnosperm species from 562 genera and 139 families. Additionally, Utkin et al. provided useful information on the SLA ranges and SLA responses to changing illumination for several species. Also, the dataset contains ~200 references to research on SLA conducted between 1918 and 2006. The dataset is ready to use in various trait analyses. There are no copyright or proprietary restrictions for research or teaching purposes. The authors would appreciate notification when and how the data are used.

Climate shapes the seed germination niche of temperate flowering plants: a meta-analysis of European seed conservation data.

BACKGROUND AND AIMS: Interactions between ecological factors and seed physiological responses during the establishment phase shape the distribution of plants. Yet, our understanding of the functions and evolution of early-life traits has been limited by the scarcity of large-scale datasets. Here, we tested the hypothesis that the germination niche of temperate plants is shaped by their climatic requirements and phylogenetic relatedness, using germination data sourced from a comprehensive seed conservation database of the European flora (ENSCOBASE). METHODS: We performed a phylogenetically informed Bayesian meta-analysis of primary data, considering 18 762 germination tests of 2418 species from laboratory experiments conducted across all European geographical regions. We tested for the interaction between species' climatic requirements and germination responses to experimental conditions including temperature, alternating temperature, light and dormancy-breaking treatments, while accounting for between-study variation related to seed sources and seed lot physiological status. KEY RESULTS: Climate was a strong predictor of germination responses. In warm and seasonally dry climates the seed germination niche includes a cold-cued germination response and an inhibition determined by alternating temperature regimes and cold stratification, while in climates with high temperature seasonality opposite responses can be observed. Germination responses to scarification and light were related to seed mass but not to climate. We also found a significant phylogenetic signal in the response of seeds to experimental conditions, providing evidence that the germination niche is phylogenetically constrained. Nevertheless, phylogenetically distant lineages exhibited common germination responses under similar climates. CONCLUSION: This is the first quantitative meta-analysis of the germination niche at a continental scale. Our findings showed that the germination niches of European plants exhibit evolutionary convergence mediated by strong pressures at the macroclimatic level. In addition, our methodological approach highlighted how large datasets generated by conservation seed banking can be valuable sources to address questions in plant macroecology and evolution.

A Compendium of in vitro Germination Media for Pollen Research.

The correct choice of in vitro pollen germination media (PGM) is crucial in basic and applied pollen research. However, the methodological gaps (e.g., strong focus of current research on model species and cultivated plants along with the lack of general rules for developing a PGM) makes experimenting with pollen difficult. We closed these gaps by compiling a compendium of optimized in vitro PGM recipes from more than 1800 articles published in English, German, and Russian from 1926 to 2019. The compendium includes 1572 PGM recipes successfully used to germinate pollen grains or produce pollen tubes in 816 species representing 412 genera and 114 families (both monocots and dicots). Among the 110 components recorded from the different PGM recipes, sucrose (89% of species), H3BO3 (77%), Ca2+ (59%), Mg2+ (44%), and K+ (39%) were the most commonly used PGM components. PGM pH was reported in 35% of all studies reviewed. Also, we identified some general rules for creating PGM for various groups of species differing in area of research (wild and cultivated species), phylogenetic relatedness (angiosperms vs. gymnosperms, dicots vs. monocots), pollen physiology (bi- and tri-cellular), biochemistry (starchy vs. starchless pollen grains), and stigma properties (dry vs. wet), and compared the component requirements. Sucrose, calcium, and magnesium concentrations were significantly different across most categories indicating that pollen sensitivity to sugar and mineral requirements in PGM is highly group-specific and should be accounted for when composing new PGM. This compendium is an important data resource on PGM and can facilitate future pollen research.

Siberian plants shift their phenology in response to climate change.

Siberia has undergone dramatic climatic changes due to global warming in recent decades. Yet, the ecological responses to these climatic changes are still poorly understood due to a lack of data. Here, we use a unique data set from the Russian 'Chronicles of Nature' network to analyse the long-term (1976-2018) phenological shifts in leaf out, flowering, fruiting and senescence of 67 common Siberian plant species. We find that Siberian boreal forest plants advanced their early season (leaf out and flowering) and mid-season (fruiting) phenology by -2.2, -0.7 and -1.6 days/decade, and delayed the onset of senescence by 1.6 days/decade during this period. These mean values, however, are subject to substantial intraspecific variability, which is partly explained by the plants' growth forms. Trees and shrubs advanced leaf out and flowering (-3.1 and -3.3. days/decade) faster than herbs (-1 day/decade), presumably due to the more direct exposure of leaf and flower buds to ambient air for the woody vegetation. For senescence, we detected a reverse pattern: stronger delays in herbs (2.1 days/decade) than in woody plants (1.0-1.2 days/decade), presumably due to the stronger effects of autumn frosts on the leaves of herbs. Interestingly, the timing of fruiting in all four growth forms advanced at similar paces, from 1.4 days/decade in shrubs to 1.7 days/decade in trees and herbs. Our findings point to a strong, yet heterogeneous, response of Siberian plant phenology to recent global warming. Furthermore, the results highlight that species- and growth form-specific differences among study species could be used to identify plants particularly at risk of decline due to their low adaptive capacity or a loss of synchronization with important interaction partners.

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.

Nikolaeva et al.'s reference book on seed dormancy and germination.

Despite the recent advances in seed science research, information on seed dormancy and germination traits is still missing for many lineages of the seed plants. We translated and digitized a huge data set on seed dormancy and germination from the Reference Book On Dormant Seed Germination by M. Nikolaeva, M. Razumova, and V. Gladkova published in Russian in 1985 and previously not readily accessible to the great majority of people in the international scientific community interested in seeds. The data set contains information on the seed dormancy classification (sensu Nikolaeva et al., 1985; Baskin and Baskin, 2004), dormancy-breaking conditions/germination, and for some of the included species storage behaviour, of seeds of nearly 3000 gymnosperm and angiosperm species from 843 genera and 164 families occurring worldwide. Additionally, the data set contains about 550 references, many of them unknown to seed scientists, to seed ecology research conducted between 1926 and 1985. The data set should be of value to applied, basic, and theoretical plant biologists, ecologists, and evolutionary biologists interested in the various aspects of regeneration of plants from seeds. There are no copyright or proprietary restrictions for research or teaching purposes. The originator of the data would appreciate notification when and how the data are used. When used in published analyses, this paper should be referred to as the data source.

Seed germination traits shape community assembly along a hydroperiod gradient.

BACKGROUND AND AIMS: Hydroperiod drives plant community composition in wetlands, resulting in distinct zonation patterns. Here, we explored the role of seed germination traits in shaping wetland community assembly along a hydroperiod gradient. Specifically, we tested the hypothesis that seeds of reed, mudflat, swamp, shallow- and deep-water communities only germinate under a specific set of environmental factors characterized by the community-specific optimal conditions for seedling survival and growth. METHODS: In a three-factorial experiment, we tested the seed germination response of 50 species typical for temperate wetlands of Europe to temperature fluctuations (constant vs. fluctuating temperature), illumination (light vs. darkness) and oxygen availability (aerobic vs. hypoxia). Phylogenetic principal component analysis, cluster analysis and phylogenetic linear regressions were used to confirm the community-specific seed germination niches. KEY RESULTS: Our study revealed the presence of five distinct, community-specific seed germination niches that reflect adaptations made by the study communities to decreasing light intensity, temperature fluctuations and oxygen availability along the hydroperiod gradient. Light as a germination trigger was found to be important in mudflats, swamps and shallow water, whereas the seeds of reed and deep-water species were able to germinate in darkness. A fluctuating temperature is only required for seed germination in mudflat species. Germination of species in the communities at the higher end of the hydroperiod gradient (reed and mudflat) demonstrated a strict requirement for oxygen, whereas swamp, shallow- and deep-water species also germinated under hypoxia. CONCLUSIONS: Our study supports the recent argument that the inclusion of seed germination traits in community ecology adds significant insights to community response to the abiotic and biotic environment. Furthermore, the close relationship between seed germination adaptations and community assembly could help reach a better understanding of the existing patterns of wetland plant distribution at local scales and wetland vegetation dynamics, as well as facilitate nature conservation measures and aquatic habitat restoration.

An Unexplored Side of Regeneration Niche: Seed Quantity and Quality Are Determined by the Effect of Temperature on Pollen Performance.

In 1977, Peter Grubb introduced the regeneration niche concept, which assumes that a plant species cannot persist if the environmental conditions are only suitable for adult plant growth and survival, but not for seed production, dispersal, germination, and seedling establishment. During the last decade, this concept has received considerable research attention as it helps to better understand community assembly, population dynamics, and plant responses to environmental changes. Yet, in its present form, it focuses too much on the post-fertilization stages of plant sexual reproduction, neglecting the fact that the environment can operate as a constraint at many points in the chain of processes necessary for successful regeneration. In this review, we draw the attention of the plant ecology research community to the pre-fertilization stages of plant sexual reproduction, an almost ignored but important aspect of the regeneration niche, and their potential consequences for successful seed production. Particularly, we focus on how temperature affects pollen performance and determines plant reproduction success by playing an important role in the temporal and spatial variations in seed quality and quantity. We also review the pollen adaptations to temperature stresses at different levels of plant organization and discuss the plasticity of the performance of pollen under changing temperature conditions. The reviewed literature demonstrates that pre-fertilization stages of seed production, particularly the extreme sensitivity of male gametophyte performance to temperature, are the key determinants of a species' regeneration niche. Thus, we suggest that previous views stating that the regeneration niche begins with the production of seeds should be modified to include the preceding stages. Lastly, we identify several gaps in pollen-related studies revealing a framework of opportunities for future research, particularly how these findings could be used in the field of plant biology and ecology.

Tree seed traits' response to monsoon climate and altitude in Indian subcontinent with particular reference to the Himalayas.

Seed traits are related to several ecological attributes of a plant species, including its distribution. While the storage physiology of desiccation-sensitive seeds has drawn considerable attention, their ecology has remained sidelined, particularly how the strong seasonality of precipitation in monsoonal climate affects their temporal and spatial distribution. We compiled data on seed mass, seed desiccation behavior, seed shedding, and germination periodicity in relation to monsoon and altitude for 198 native tree species of Indian Himalayas and adjoining plains to find out (1) the adaptive significance of seed mass and seed desiccation behavior in relation to monsoon and (2) the pattern of change in seed mass in relation to altitude, habitat moisture, and succession. The tree species fall into three categories with respect to seed shedding and germination periodicities: (1) species in which both seed shedding and germination are synchronized with monsoon, referred to as monsoon-synchronized (MS, 46 species); (2) species in which seed germination is synchronized with monsoon, but seeds are shed several months before monsoon, referred to as partially monsoon-synchronized (PMS, 112 species); and (3) species in which both shedding and germination occur outside of monsoon months, referred to as monsoon-desynchronized (MD, 39 species). The seed mass of MS species (1,718 mg/seed) was greater than that of PMS (627 mg/seed) and MD (1,144 mg/seed). Of the 40 species with desiccation-sensitive seeds, 45% belong to the MS category, almost similar (approx. 47%) to woody plants with desiccation-sensitive seeds in evergreen rain forests. Seed mass differed significantly as per seed desiccation behavior and successional stage. No relationship of seed mass was found with altitude alone and on the basis of seed desiccation behavior. However, seed mass trend along the altitude differed among monsoon synchronization strategies. Based on our findings, we conclude that in the predicted climate change (warming and uncertain precipitation pattern) scenario, a delay or prolonged break-spell of monsoon may adversely affect the regeneration ecology of desiccation-sensitive seed-bearing species dominant over large forest areas of monsoonal climate.

Contrasting Effects of Extreme Drought and Snowmelt Patterns on Mountain Plants along an Elevation Gradient.

Despite the evidence that increased frequency and magnitude of extreme climate events (ECE) considerably affect plant performance, there is still a lack of knowledge about how these events affect mountain plant biodiversity and mountain ecosystem functioning. Here, we assessed the short-term (one vegetation period) effects of simulated ECEs [extreme drought (DR), advanced and delayed snowmelt (AD and DE), respectively] on the performance of 42 plant species occurring in the Bavarian Alps (Germany) along an elevational gradient of 600-2000 m a.s.l. in terms of vegetative growth and reproduction performance. We demonstrate that plant vegetative and generative traits respond differently to the simulated ECEs, but the nature and magnitude treatment effects strongly depend on study site location along the elevational gradient, species' altitudinal origin and plant functional type (PFT) of the target species. For example, the negative effect of DR treatment on growth (e.g., lower growth rates and lower leaf nitrogen content) and reproduction (e.g., lower seed mass) was much stronger in upland sites, as compared to lowlands. Species' response to the treatments also differed according to their altitudinal origin. Specifically, upland species responded negatively to extreme DR (e.g., lower growth rates and lower leaf carbon concentrations, smaller seed set), whereas performance of lowland species remained unaffected (e.g., stable seed set and seed size) or even positively responded (e.g., higher growth rates) to that treatment. Furthermore, we were able to detect some consistent differences in responses to the ECEs among three PFTs (forbs, graminoids, and legumes). For instance, vegetative growth and sexual reproduction of highly adaptable opportunistic graminoids positively responded to nearly all ECEs, likely on the costs of other, more conservative, forbs and legumes. Our results suggest that ECEs can significantly modify the performance of specific plant groups and therefore lead to changes in plant community structure and composition under ongoing climate change. Our study therefore underlines the need for more experimental studies on the effects of extreme climate events to understand the potential consequences of climate change for the alpine ecosystem.

Effects of extreme drought on specific leaf area of grassland species: A meta-analysis of experimental studies in temperate and sub-Mediterranean systems.

Here, we conducted a meta-analysis of experimental drought manipulation studies using rainout shelters in five sites of natural grassland ecosystems of Europe. The single studies assess the effects of extreme drought on the intraspecific variation of the specific leaf area (SLA), a proxy of plant growth. We evaluate and compare the effect size of the SLA response for the functional groups of forbs and grasses in temperate and sub-Mediterranean systems. We hypothesized that the functional groups of grasses and forbs from temperate grassland systems have different strategies in short-term drought response, measured as adjustment of SLA, with SLA-reduction in grasses and SLA-maintenance in forbs. Second, we hypothesized that grasses and forbs from sub-Mediterranean systems do not differ in their drought response as both groups maintain their SLA. We found a significant decrease of SLA in grasses of the temperate systems in response to drought while SLA of forbs showed no significant response. Lower SLA is associated with enhanced water-use efficiency under water stress and thus can be seen as a strategy of phenotypic adjustment. By contrast, in the sub-Mediterranean systems, grasses significantly increased their SLA in the drought treatment. This result points towards a better growth performance of these grasses, which is most likely related to their strategy to allocate resources to belowground parts. The observed SLA reduction of forbs is most likely a direct drought response given that competitive effect of grasses is unlikely due to the scanty vegetation cover. We point out that phenotypic adjustment is an important driver of short-term functional plant response to climatic extremes such as drought. Differential reactions of functional groups have to be interpreted against the background of the group's evolutionary configuration that can differ between climatic zones.

Minimal temperature of pollen germination controls species distribution along a temperature gradient.

BACKGROUND AND AIMS: Although plant distribution patterns are well documented, our understanding of the ecophysiological mechanisms that control the geographical ranges of plant species remains poor. We used a largely ignored method, the performance of the male gametophyte in vitro, to assess whether the thermal range of pollen germination and tube growth controls species distribution ranges, in this case along an elevational gradient. METHODS: Using in vitro pollen germination experiments, we obtained cardinal temperatures (minimal, optimal and maximal) of pollen germination and pollen tube growth for 25 herbaceous species along a mean annual temperature gradient of about 5 °C. These temperatures were correlated with temperatures of the sites where the species were collected. The presence of a phylogenetic signal in the data set as well as an effect of species flowering phenology were also estimated. KEY RESULTS AND CONCLUSIONS: We found a strong positive relationship between temperature conditions at our collection sites and the minimum temperature for both pollen germination and pollen tube growth. In addition, a significant correlation between maximum temperature of pollen tube growth and temperature of flowering month was apparent. We conclude that the restriction of pollen germination and growth by low temperatures is an important contributor to the climatic restriction of plant species distributions. Improved knowledge of this thermal precursor to seed production could, from a functional perspective, enhance our understanding of species distributions along climatic gradients and our ability to predict how anthropogenic climate change might affect plant community composition.