Can germination explain the distribution of tree species in a savanna wetland?

The relationship between seed germination and ecological niche is determined by matching germination characteristics with environmental features. In this study, we selected tree species occurring in the largest savanna wetland in South America – the Pantanal. Very few species are endemic or exclusively found in savanna wetlands, and the majority of tree species occurring in the Pantanal are also found in the neighbouring Brazilian Cerrado, a drier vegetation type that does not flood. We investigated the relationship between germination characteristics and occurrence of savanna trees in wetlands testing the hypothesis that such seeds are tolerant to flooding. We also addressed the question of whether seed tolerance to flood, assessed by survival analysis, explains tree distribution along a gradient of flooding intensity. In this flooding gradient, widely distributed species are those that occur in areas subjected to low as well as to high flooding intensity whereas restricted distributed species are those that occur only in areas subjected to a low level of flood. Seeds from tree species occurring in areas subjected to different flooding intensities were collected. Seed tolerance and germination during and after both one and two months of simulated flood were evaluated. Our results show that seeds of most of the studied savanna species tolerated submergence, which helps to explain their occurrence and wide distribution in wetlands. Nevertheless, germination behaviour checked by survival functions (i.e. how germination is distributed over time) partially explained tree species distribution along a flooding gradient. We conclude that seed tolerance to flooding is one of the components of the regeneration niche that determines tree occurrence and distribution at the regional scale, from savanna to wetland, but not at a local scale along a flooding gradient.     

Correspondence of seed traits with niche position in glacier foreland succession

Changes in species composition during succession are driven by biotic and abiotic factors leading to a multitude of niches occupied by distinct species. Gradient analyses of plant communities provide opportunities to approximate the niche position of species along a successional gradient. Several plant traits have been used to explain mechanisms governing successional sequences, but generalising changes in species traits during primary succession is still controversial. This study examined whether the seed mass and the optimum temperature for germination could explain the niche position of several glacier foreland species along a primary successional gradient in the Austrian Central Alps. We hypothesised that pioneer species should possess lighter seeds and a lower optimum temperature for germination than late successional species. We found significant differences in the seed mass between species, but the seed mass did not correspond with the assigned niche position on the successional gradient. Germination responses to temperature also differed significantly between species. Pioneer species performed better at lower temperatures than late successional species, suggesting that the optimum temperature for germination is a driver of niche separation. We discuss the interactions between seed traits and environmental conditions along the primary successional gradient emphasising the importance of temperature requirements for the germination. Differences in the regeneration characteristics are a major cue governing species turnover in glacier foreland succession.     

Seed mass and dormancy of annual plant populations and communities decreases with aridity and rainfall predictability

Several theoretical and empirical studies have examined the influence of environmental conditions on seed traits and germination strategies of annual species. A positive relationship between seed mass and dormancy has been described for annuals occupying climatically unpredictable ecosystems. Larger-seeded species tend to have higher seedling survival rates, while dormancy allows a bet-hedging strategy in unpredictable environments. Until now, these ideas have been addressed primarily for only one or a few focal species, without considering differences among populations and communities. The novelty of the present study lies in the population and community-level approach, where a comprehensive seed trait database including 158 annual species occurring along a gradient of rainfall variability and aridity in Israel was used to ask the following question: Does average seed mass and dormancy of annual populations and communities decrease with increasing aridity and rainfall unpredictability?Soil seed bank samples were collected at the end of the summer drought, before the onset of the rains, from four plant communities. Germination was tested under irrigated conditions during three consecutive germination seasons to determine the overall seed germinability in each soil sample. Seed mass was obtained from newly produced seeds collected at the study sites in late spring. The community level results showed that, in contrast to common theoretical knowledge, seed mass and dormancy of the dominant annual species decreased with increasing aridity and rainfall variability. Accordingly, a negative correlation was found between seed mass and seed germination fractions. The present study demonstrates that an analysis of seed traits along climatic gradients is significantly improved by approaches that target both population and community levels simultaneously. A critical evaluation sheds new light upon the selective pressures that act on seed ecology of annuals along a climatic gradient and facilitates formulation of more mechanistic hypotheses about factors governing critical seed traits.     

Germination of <i>Eryngium regnellii</i>: a major species for ecological restoration of plant-pollinator interactions in the Southern Pampas (Buenos Aires,Argentina)

Eryngium regnellii Malme belongs to the largest genera in the Apiaceae family, with 250 species worldwide and 65 represented in South America. It is a herbaceous species typical of hill plant communities, which, along with remnant grassland patches, are the most relevant natural habitats for the maintenance of diversity in the Southern Pampas. Eryngium regnellii is key to the maintenance of pollination mutualisms, being a generalist (displaying a diverse assemblage of pollinators) and ubiquitous species (present in all studied sierras). However, fragmentation of the Pampean landscape due to agricultural intensification has led to the loss of natural environments. Therefore, the reintroduction of E. regnellii in strategic places would facilitate the occurrence of wild pollinators, while favoring pollination services in the agroecosystem. The germination requirements of E. regnellii were studied because a better knowledge of the reproductive biology of this species would provide information relevant to its reproduction and reintroduction into degraded areas. Germination percentages and mean time to germination were evaluated, using one control and two pre-germination treatments: chemical scarification with sulfuric acid, and mechanical scarification with sand paper. Chemical scarified seeds did not germinate. Mechanically scarified and control seed groups showed no significant differences either in germination percentages (49% and 59% respectively) or in mean germination time (13 and 14 days, respectively). Results indicate that E. regnellii shows no physical dormancy, and does not require specific pre-germination treatments for germination under the studied laboratory conditions. The high germination capacity of E. regnellii, along with its ecological attributes, make it a potential species for restoring plant-pollinator interactions in the fragmented landscapes of the Southern Pampas.     

Interspecific and intraspecific variation in seed size and germination requirements of Sarracenia (Sarraceniaceae)

Seed size and germination requirements of eight (of nine) Sarracenia species, and 13 populations of S. purpurea were studied. All species except for S. purpurea are restricted to the southeastern United States, whereas S. purpurea ranges across Canada, southward along the eastern United States into Maryland and Virginia (S. purpurea ssp. purpurea), and from New Jersey southward into northern Florida and the coast of the Gulf of Mexico (S. purpurea ssp. venosa). I tested the hypotheses that dormancy-breaking requirements vary predictably among species across a latitudinal gradient. I also sought to determine whether seed size and germination requirements were useful characters for resolving systematic and phylogenetic questions within this genus. Seed size varied significantly among species, but variability in seed size within S. purpurea exceeded the variability in seed size observed across all eight species studied. Seeds of all species are morphophysiologically dormant upon dispersal. Length of required cool, moist pretreatment varied among species, and germination in higher latitude populations is enhanced with longer pretreatment. In contrast, variability in germination requirements of subspecies, varieties, and populations of the geographically wide-ranging S. purpurea was not related clearly to geographic location (latitude or elevation). Germination requirements do not map onto a proposed phylogeny of Sarracenia, but observed differences in germination requirements of S. purpurea ssp. venosa var. burkii relative to other populations of S. purpurea support the recent proposal to elevate this variety to species status.     

Impact of fire on Picea mariana and Pinus banksiana seedlings in subarctic lichen woodlands

Abstract. The demography of Picea mariana (black spruce) and Pinus banksiana (jack pine) seedlings was monitored through five censuses over 13 months in four different seed bed types after fire of four severity levels in lichen woodland. Most seeds germinated just before early frost in late summer 1990 or immediately after snow thaw in early spring 1991; the germination rate subsequently decreased. For both species, germination rate decreased along a gradient of fire severity. The proportion of Pinus seeds that produced a seedling surviving 13 months after sowing was 4.3 % and 0.4 % respectively in the intact lichen mat and in the mineral soil seed bed type. For Picea these values are 3.2 % and 0.2 % respectively. The low germination rate in a severely burned seed bed type appeared to be associated with the formation of a water-repellent crust at the soil surface following the fire. Seedlings were contagiously distributed and were more frequent in flat and hollow microsites, where there is probably more water available than on bumps or among pebbles. Properties of experimentally burned seed bed types may differ from those under natural fires where regeneration by seed generally occurs following dispersal. However, the high germination rate observed in the intact lichen mat suggests that scattered lichen woodland patches may respond to increased seed input by a higher frequency of seedling establishment.     

The effect of water supply on seed-bank analysis using the seedling-emergence method

1. When using the seedling-emergence method to analyse a soil seed bank the greenhouse conditions should match the germination requirements of the species involved. Although the seedling-emergence method is common practice in ecological studies, the germination characteristics of many species are not known, or are only partly known.
2. Before carrying out a large-scale seed-bank study in a wet dune slack, we tested the water requirements of the species in the seed bank, comparing a waterlogged soil with a moist soil. Four species germinated in significantly larger numbers in a waterlogged soil, seven species in a moist soil and 11 species showed no significant difference. When a species was present in low densities, it was often missed using the less appropriate treatment.
3. These results emphasize the need for a preliminary study before carrying out a seed-bank analysis and show the danger of using a 'standard' method to analyse soil samples from different habitats.     

Seed bank of seasonally flooded grassland: experimental simulation of flood and post-flood

Wetland seed banks comprise the propagules of plant species that have species-specific germination requirements for germination in either flooded or dry conditions. At the community level, wetland structure and succession during and after a seasonal flooding event depends upon the early life-history requirements of species, including germination under flooded and dry conditions. We examined the effects of simulated flood and post-flood scenarios on seedling emergence from a seed bank of seasonally flooded grassland in the Pantanal, Brazil. Field samplings were conducted in both wet and dry seasons, both of which were subject to flood and post-flood conditions. A total of 70 species emerged from the seed bank, dominated by Poaceae and Cyperaceae. Sixteen species were exclusive to the wet and one exclusive to the dry season. The richness of perennial species was higher under flood conditions, while the richness of annuals was greater post-flood. In general, the aquatic and amphibious species exhibited a significant germination response to flooding. Terrestrial species only germinated in post-flood conditions, with higher richness in the dry season. Four species had high seedling abundance in both treatments. The capacity of regeneration by seeds is high in these grasslands and can be increased by seasonal flooding and drawdown. In these seasonally flooded grasslands, we observed three main germination strategies: under flooded conditions, aquatic and amphibious species; post-flood conditions, an explosion of annual amphibious and terrestrial species; and in moist soil, perennial terrestrial species. The differential responses to flooding versus post-flood conditions help to maintain the structure and species richness in the community over time.     

短命植物独行菜种子萌发过程对低温的耐受特性

以独行菜（Lepidium apetalum）为材料,研究了其种子在萌发过程中耐受低温的特性,并对耐受低温的机制作了初步的探讨。结果表明：1.萌发至I、II、III期的独行菜种子经过冷诱导处理后,对零下低温-5℃、-10℃胁迫具有较强的耐受性,这种耐受性随萌发发展相对有所降低。2.独行菜种子萌发过程中胚根生长速度明显受到低温抑制,但也仍然能够生长。3.适当时间的低温层积能使独行菜萌发势显著提高,对终萌发率影响不大;过长时间的低温层积会使独行菜种子萌发势和最终萌发率降低。4.独行菜种子在4℃条件下不能萌发,但4℃层积2d的种子经25℃处理6h以上后,可耐受4℃低温而萌发,且萌发率随25℃处理时间增加而升高。如果25℃处理时间少于5h则不能在4℃低温下萌发。经4℃层积10d、再经25℃萌发处理1h的独行菜种子就能在4℃低温下萌发。综合分析认为独行菜种子不能耐受4℃低温萌发,原因可能是在露白前存在一个关键的生理阶段,在4℃胁迫逆境中不能越过这个阶段,该阶段之前与之后的萌发过程都能耐受4℃低温,因此对低温胁迫有良好的耐受性。这为探索早春短命植物耐受低温萌发的机制提供了新的思路。     

Divergence in germination traits among arctic and alpine populations of <Emphasis Type="Italic">Koenigia islandica</Emphasis>: light requirements

Light is known to regulate conservative germination strategies and the formation of seed banks. Although these strategies are crucial to survival in tundra environments—especially for annuals—light requirements for germination in arctic–alpine species are seldom investigated. Furthermore, environmental differences between arctic and alpine regions are expected to lead to evolutionary divergence among conspecific populations in seed germination strategies. In this study, we report important differences in germination light requirements among six arctic and alpine populations of the annual Koenigia islandica. Light had little effect on germination of the seeds from Iqaluit (Nunavut, Canada), Yukon (Canada), and Jasper (Alberta, Canada), whereas the seeds from the most severe climates, Svalbard (Norway) and Colorado (USA), had strong light requirements. Stratification of the seeds had little influence on their germination light requirements, with the exception of the population from Dovre (Norway), in which it induced a strong light requirement. Possible adaptive explanations and some implications of these observed germination patterns are discussed.     

Indirect effect of abscisic acid on the induction of secondary dormancy in lettuce seeds

During temporary incubation at 25°C in buffered solutions (pH 4.0) of abscisic acid (ABA) seeds of lettuce ( Lactuca sativa L. cv. Olof) lost the red-light initiated ability to germinate in buffer. The development of secondary dormancy required an inhibitory ABA content in the seeds during a number of days. A temporary incubation in ABA during 24 h met these requirements only if the solution was about 100-fold more concentrated than during continuous incubation. Studies with 2-¹⁴C-ABA showed that the amount of ABA which had penetrated in 24 h was reduced by a factor 100 within 3 to 4 days during subsequent incubation in buffer. Both leaching and metabolic changes were involved in the reduction process. The nature of the metabolic products remained obscure. A shift to 2°C after incubation in ABA prevented the induction of secondary dormancy, but inhibited ABA metabolism. ABA did not interfere with the induction rate of secondary dormancy, and it was not required to maintain the state of dormancy. The sole function of ABA was the non-specific inhibition of germination, which indirectly facilitated the development of an ABA independent secondary dormancy. – The level of endogenous ABA was compared to the amount of ABA found in the embryo during and after incubation in ABA solutions marked with 2-¹⁴C-ABA. The level of endogenous ABA in air-dry seeds (0.11 ng/mg dry weight) corresponded to the minimal level at which penetrated ABA inhibited germination. This level had to be present at least during 4 to 5 days to inhibit the effect of red light. Since endogenous ABA was quickly reduced upon imbibition, a regulatory function of endogenous ABA in the inhibition of red light induced germination can be ruled out. A function in the temporary inhibition of dark germination and, consequently, in the development of secondary light irresponsiveness cannot be excluded, however.     

Leaf gas films,underwater photosynthesis and plant species distributions in a flood gradient

Traits for survival during flooding of terrestrial plants include stimulation or inhibition of shoot elongation, aerenchyma formation and efficient gas exchange. Leaf gas films form on superhydrophobic cuticles during submergence and enhance underwater gas exchange. The main hypothesis tested was that the presence of leaf gas films influences the distribution of plant species along a natural flood gradient. We conducted laboratory experiments and field observations on species distributed along a natural flood gradient. We measured presence or absence of leaf gas films and specific leaf area of 95 species. We also measured, gas film retention time during submergence and underwater net photosynthesis and dark respiration of 25 target species. The presence of a leaf gas film was inversely correlated to flood frequency and duration and reached a maximum value of 80% of the species in the rarely flooded locations. This relationship was primarily driven by grasses that all, independently of their field location along the flood gradient, possess gas films when submerged. Although the present study and earlier experiments have shown that leaf gas films enhance gas exchange of submerged plants, the ability of species to form leaf gas films did not show the hypothesized relationship with species composition along the flood gradient.     

香圆种子萌发特征及催芽技术

以香圆（Citrus wilsonii Tanaka）种子为实验材料,通过测定种皮透水性、种皮机械障碍、发芽温度等对种子萌发的影响从而探讨种子萌发特征;通过切割种子、热水浸种、赤霉素浸种、低温层积等处理方式探索种子的催芽方法。结果显示：香圆种皮不存在透水性障碍,但其机械障碍对种子萌发具有抑制作用,且内种皮对萌发影响较大;种子在15℃、25℃、35℃恒温及15℃/30℃变温条件下均可萌发,其最适萌发温度为25℃;采用低温层积、热水浸种、切除1/3种子等方法均可一定程度上促进种子萌发。生产中宜采用始温60℃热水浸种24h的方法对香圆种子进行催芽。     

干藏和淹水对三峡库区21种草本植物种子萌发的影响

研究了三峡库区河岸带21种常见草本植物种子在干藏以及不同淹水深度（0.5、1.0和2.0m）处理下的萌发特性。结果表明：以新采集种子的萌发率作为对照,干藏后8个物种的萌发率增加（占总数的38.1%）,1种萌发率不变（4.8%）,12种萌发率降低（57.1%）。与对照相比,淹水后3个物种的萌发率增加（占总数的14.3%）,2个物种的萌发率不变（9.5%）,16个物种的萌发率下降（76.2%）。水淹处理后,占总数76.2%的物种的萌发率下降,其中11种显著下降;但淹水深度对种子的萌发没有显著影响。干藏后的繁穗苋（Amaranthus paniculatus）、皱果苋（Amaranthus viridis）和红刺果苋（Amaranthus sp.）以及水淹后的鳢肠（Eclipta prostrata）萌发率〉50%,而且能够利用库区退水期完成生活史形成种子,表明这些植物可作为三峡库区消落带植被恢复时优先考虑的物种源。     

Temperature requirements for seed germination of Pereskia aculeata and Pereskia grandifolia

Pereskia aculeata and Pereskia grandifolia have been studied widely due to their high nutritional and therapeutic values. However, little is known about the biological requirements of their seeds for the various germination factors. Thus, this experiment aimed to evaluate the thermal effects on the germination of these species at the temperatures of 24 °C, 27 °C, 30 °C, 33 °C and 36 °C. After verification of the existence of differences in the performance of germination, a non-linear regression was carried out, relating the germination to temperature and identifying its point of maximum efficiency. We found that the lowest synchronization indexes of germination were observed close to 30 °C. The best germination response of the P. aculeata and P. grandifolia was observed at 30 °C and 33 °C, respectively, with greater germination strength and fewer days to attain 63.21% of germinations. The results obtained from the germination of P. aculeata and P. grandifolia can be described by the Weindull distribution model with three parameters, as proposed by Carneiro and Guedes (1992).     

The Characteristics of Low Temperature Tolerance during Seed Germination of the Ephemeral Plant Lepidium apetalum (Cruciferae)

Lepidium apetalum was used to explore the characteristics and mechanismes of low temperature tolerance during seed germination. The results showed that pre treatment the seeds at 4℃ would improve the seeds viability after stressed by low temperature at -5℃ and -10℃ and the capability of seeds tolerance to cold temperature became weaker along with germination development. Although the growth of radicle was inhibited significantly at 4℃ during germination, the seeds could still survive and grow. With appropriate time of stratified at 4℃, germination pace would be improved and germination rate would not change, while both of which decreased if stratification lasted too long. Moreover, Lapetalum seeds were very sensible to the temperature during germination. It showed no germination at 4℃, while if seeds were stratificated for 48 hours at 4℃ and subsequently treated with 25℃ for more than 6 hours, seeds would germinate at 4℃ and the germination rate increased with the time of 25℃ treatment prolonged. If treatment at 25℃ was less than 5 houes, it showed no germinating. In addition, seeds with stratification for 10 days following 1 h treatment at 25℃ could also germinated at 4℃. The present study indicated that some pivotal genes that related with a key physiological stage before the onset of radicle protrusion might inhibite at 4℃, which resulted in Lapetalum seed could not germinate at 4℃. Except that physiological stage, the Lapetalum seed had a strong tolerance to low temperature at other germination stages. This experiment provided a new way to explore the germination mechanism of ephemerals plants seeds at low temperature.     

三峡濒危植物疏花水柏枝的回归引种和种群重建

疏花水柏枝分布于三峡库区原海拔70～155m的消落带,三峡工程修建后它将丧失其全部生境而成为濒危植物。实验结果显示其种子在土壤含水量大于10%以上时开始萌发,以土壤含水量达到饱和状况时萌发最好。种子萌发与定居阶段对土壤水分条件的严格要求使得疏花水柏枝分布区十分狭小。回归引种和种群重建是拯救该物种的主要手段。三峡工程修建后库区内新的消落带将形成夏旱冬淹的水节律,完全不同于库区原有消落带所具有的冬旱夏淹的水节律,不适于作为疏花水柏枝种群的迁移地。相比之下库区淹没区以上各支流消落带的生态环境与疏花水柏枝原有生境较为接近,适于作其新的生境。种群遗传多样性、年龄结构、分布格局、繁殖与扩展等生物学特性是种群持续发展的基础,文章以此为依据,对疏花水柏枝种群重建与管理中的相关问题进行了分析讨论。认为疏花水柏枝种群恢复与重建中目前所面临的主要问题是如何增强被隔离的种群间的基因交流、促进种群的种子扩散与萌发、协调新建种群与当地物种的关系、营造有利于新建种群定居与生长的生态环境。重建种群的管理应结合疏花水柏枝的生长发育节律和移栽地的生态环境条件来开展,要有效地监控种群的生长发育动态,合理地在隔离种群间相互引种,适时地进行水分管理,并对周围植被适度控制。     

渗透胁迫下水稻种子萌发特性及抗旱性鉴定指标研究

用聚乙二醇（PEG）-6000作为渗透剂模拟干旱胁迫,探讨在4种不同浓度的渗透胁迫下水稻种子的萌发特性,并确立间接评价水稻苗期抗旱性的萌发期鉴定指标。结果表明,高浓度渗透液对种子萌发的抑制作用明显大于低浓度,萌发率的干旱胁迫反应指数随渗透液浓度的降低和萌发进程而增加。在不同浓度的渗透胁迫处理中,20％浓度下萌发6d后的萌发率表现较大的变异幅度和变异系数以及较均衡的品种频率分布,较好地反映了水稻种质对渗透胁迫的不同反应,认为20％／6d条件较适合作为萌发率鉴定的渗透液模拟干旱胁迫;经反复干旱后的幼苗存活率与萌发胁迫指数和20％胁迫浓度下萌发8d后芽鞘长的干旱胁迫反应指数（DRI）表现极显著或显著正相关,认为萌发胁迫指数和芽鞘长的DRI可作为评价水稻苗期抗旱性的间接鉴定指标。     

The contribution of persistent soil seed banks and flooding to the restoration of alluvial meadows

Restoration of species-rich flood meadows impoverished by agricultural intensification is an important challenge. The relationships between flooding regime and soil seed bank were compared in three successive meadow communities (hygrophilic, mesohygrophilic and mesophilic) distinguished along a topographic and hydric gradient. Differences in flood duration and frequency between the three associations allowed the study of the contribution of floods to soil seed bank richness and density. No significant difference was found in species richness among the three soil seed banks, whereas the densities were significantly higher in the wettest community. The three seed bank compositions were clearly distinguished along the hydric gradient. In fact, the three seed bank types constituted a species poor version of the meadow communities to which they belong. Flood contributions appear to play a minor role in seed bank enrichment. Thus, seed dispersal by flood water would probably be insufficient to enable the restoration of alluvial meadows.     

Germination and anaerobic metabolism of seeds of Tabebuia cassinoides (Lam.) DC subjected to flooding and anoxia

Tabebuia cassinoides (Lam.) DC (Bignoniaceae) is an arboreal species common in seasonally or permanently waterlogged areas of the “restinga” forest (a type of forest that occurs on the sandbanks of the coastal plains of southeastern Brazil). The objectives of the present study were to establish seed germination responses of this species to flooding and anoxia and investigate the end products of the anaerobic metabolism of seeds subjected to these conditions, with the goal of understanding the adaptive strategies that enable this species to dominate flood prone areas of “restinga”, as well as determine reserves stored in their seeds. Seeds of T. cassinoides did not germinate under anoxia or complete submergence, but remained viable under these conditions for 15 and 20 days, respectively. Due to their membranaceous wings, the seeds float very well and reached 100% germination in this condition, an important adaptation to overcome the initial stages of development in flooded habitats. In relation to anaerobic metabolism, ethanol is the most important end product, while lactate is produced in lower concentrations. Seeds of T. cassinoides have very little endosperm and the reserves, mainly glycoproteins, lipids and free sugars, accumulate in the cotyledons. Free sugars may provide the substrate for the initial metabolism of seed germination, but the level of these reserves was not enough to allow germination under oxygen deprivation. Possibly, carbohydrate reserves were enough only to maintain seed viability for a relative short period under this condition.