A convenient screening test system and a model for thermal germination responses of wild plant seeds: behaviour of model and real seeds in the system

Abstract A convenient test system for screening the thermal germination behaviour of seeds was developed for both basic and applied research in seed germination ecophysiology. Only two temperature-controlled facilities, a test period of about a month and a relatively small number of sample seeds arc needed to obtain information on the thermal-germination parameters of individual seed populations, such as lower or higher limit temperatures, and thermal times required for germination. In the test system, the germination performances of sample seed populations were compared under two temperature regimes: a gradually increasing temperature regime and a gradually decreasing temperature regime, in which the seeds were subjected to gradually changing temperatures in the range of 4 36°C. In order to assess the effects of various values for thermal-germination parameters on the patterns of germination performance in the system, the behaviour of model seeds characterized by a definite set of thermal germination parameters were investigated. Referring to the results of the simulations, the actual germination patterns of some wild-seed populations in the test system were interpreted in terms of thermal-germination parameters.     

Differentiation in seed germination among populations of Capsicum annuum along a latitudinal gradient in Mexico

Toinvestigate variation in germination capability of wild Capsicumannuum seeds, the effects of light, fluctuating temperature,gibberellic acid (GA), soaking and sulfuric acid were analyzed in samples from14 populations from northwest Mexico. Germination was inhibited by darkness inall populations studied. Darkness inhibition of germination was overcome byfluctuating temperature and GA, factors which also increased germinationpercentage but not germination rate. Soaking treatments did not significantlyincrease germination of C. annuum seeds, while sulfuricacid treatments had negative effects on the process. Populations showed highvariation in germination response in all treatments, except for continuousdarkness. Principal component analysis differentiated some populations withhighgermination percentage and slow germination rate. Such differences did notcorrelate with climatic variables from collection sites. Given the variation ingermination patterns observed, some of these totally contrasting, we have to becautious when considering the results of studies based on a single wildpopulation.     

Rainfall seasonality predicts the germination behavior of a tropical dry‐forest vine

Seed dormancy is considered to be an adaptive strategy in seasonal and/or unpredictable environments because it prevents germination during climatically favorable periods that are too short for seedling establishment. Tropical dry forests are seasonal environments where seed dormancy may play an important role in plant resilience and resistance to changing precipitation patterns. We studied the germination behavior of seeds from six populations of the Neotropical vine Dalechampia scandens (Euphorbiaceae) originating from environments of contrasting rainfall seasonality. Seeds produced by second greenhouse‐generation plants were measured and exposed to a favorable wet environment at different time intervals after capsule dehiscence and seed dispersal. We recorded the success and the timing of germination. All populations produced at least some dormant seeds, but seeds of populations originating from more seasonal environments required longer periods of after‐ripening before germinating. Within populations, larger seeds tended to require longer after‐ripening periods than did smaller seeds. These results indicate among‐population genetic differences in germination behavior and suggest that these populations are adapted to local environmental conditions. They also suggest that seed size may influence germination timing within populations. Ongoing changes in seasonality patterns in tropical dry forests may impose strong selection on these traits.     

A spatially explicit analysis of seedling recruitment in the terrestrial orchid Orchis purpurea

Seed dispersal and the subsequent recruitment of new individuals into a population are important processes affecting the population dynamics, genetic diversity and spatial genetic structure of plant populations. Spatial patterns of seedling recruitment were investigated in two populations of the terrestrial orchid Orchis purpurea using both univariate and bivariate point pattern analysis, parentage analysis and seed germination experiments. Both adults and recruits showed a clustered spatial distribution with cluster radii of c. 4-5 m. The parentage analysis resulted in offspring-dispersal distances that were slightly larger than distances obtained from the point pattern analyses. The suitability of microsites for germination differed among sites, with strong constraints in one site and almost no constraints in the other. These results provide a clear and coherent picture of recruitment patterns in a tuberous, perennial orchid. Seed dispersal is limited to a few metres from the mother plant, whereas the availability of suitable germination conditions may vary strongly from one site to the next. Because of a time lag of 3-4 yr between seed dispersal and actual recruitment, and irregular flowering and fruiting patterns of adult plants, interpretation of recruitment patterns using point patterns analyses ideally should take into account the demographic properties of orchid populations.     

古尔班通古特沙漠及周缘52种植物种子的萌发特性与生态意义

本文对古尔班通古特沙漠及其周缘52种植物种子的萌发特性进行比较,以探讨该地区植物的不同萌发对策及其生态适应意义。结果发现：萌发率80%以上的有11种,小于20%的有22种,7种未萌发;萌发开始时间为1-3d的有37种,大于10d的有2种;萌发持续时间为1-7d的有10种,大于22d的有14种;达50%萌发率的时间为1-7d的有17种。主成分分析(PCA)和聚类分析结果表明,52种植物可划分为4种萌发类型：爆发型、过渡型、缓萌型和低萌型,这可能与生境土壤中水分存在差异有关;木本植物的萌发率显著高于非木本植物,暗示木本植物的高萌发率保证植物能够快速占领空间和资源,增加竞争优势,而非木本植物,尤其是一年生植物表现低的萌发率,是植物生存的一种风险分摊策略。     

Climatic control of germination in the genus Silene L

Background: Seed germination strategies depend on a number of factors such as climatic variables, habitat, life cycle, phylogenetic affiliation and morphological seed traits.

Aims: We present a novel evaluation of the ecological correlates of germination strategies focussing on plant populations drawn from a single genus and considering a number of relevant explanatory variables.

Methods: Temperature and light-controlled germination experiments were carried out on 21 seed accessions belonging to 13 closely related species of the genus Silene L. Accessions were selected to tightly control the phylogenetic effect and give broad ecological and geographic coverage of the genus in Europe, with three North American and Macaronesian outgroups from climatically extreme environments. We used principal component analysis and correlation analyses to identify the correlation structure of germination traits and include multiple explanatory variables.

Results: Three germination strategies were shown to be related to climatic control. (1) Seeds from arid regions with hot, dry summers germinated optimally at cool temperatures associated with the rainfall period under field conditions, whereas (2) the benefit of cold stratification was more marked in provenances characterised by cold, dry winters, and (3) seeds from mild climates preferred warm temperatures for germination. Moreover, (4) biennial populations of disturbed habitats showed synchronised and rapid seed germination over a wide thermal window. In agreement with previous findings, (5) habitat-related syndromes were observed only for wetland populations. Correlations with seed mass were significant when related to summer precipitation, but weak or absent in relation to germination traits, indicating that, though influenced by the local climate, seed mass is a poor predictor of germination strategies.

Conclusions: These results suggest that whilst habitat and life cycle might shape germination patterns to a certain extent, long-term climatic differences play a substantial role in selecting specific germination traits and strategies.     

Effect of storage time,site and floral morph on seed germination of the threatened distylous primrose Primula cusickiana var. maguirei

Ex situ conservation of rare plant species requires an understanding of germination requirements. We report the first successful germination trials for Maguire primrose (Primula cusickiana var. maguirei[L.O. Williams] N.H. Holmgren & S. Kelso), a threatened perennial herb narrowly restricted to a 20‐km canyon in northern Utah, USA. Seeds collected from different sites (upper or lower canyon populations) and flower morphs (pin or thrum) that had been stored for either 1 or 2 years were cold, moist stratified in darkness at 1–2°C for 270 days. Independent of treatment, time to first germination was approximately 75 days, time to 50% germination (T₅₀) was approximately 165 days, and total germination was 55.8%. Higher mean germination was observed in seeds stored for 2 years compared with seeds stored for 1 year, as well as in seeds from pin flowers compared with thrum flowers. There was not a significant effect of site on mean germination, but there was a significant storage time × site interaction. Seeds stored for 2 years also had a lower mean T₅₀ versus seeds stored for 1 year. In addition, seeds from upper canyon populations had a higher mean T₅₀ than those from lower canyon populations. Our results suggest that short‐term storage should not significantly decrease seed viability, and that both flower morph and canyon location may influence germination. Evidence suggests that germination patterns are adaptive. Our results contribute to our understanding of the germination biology of P. cusickiana var. maguirei and inform future conservation efforts for this threatened species.     

Changes in mRNA populations during loblolly pine (Pinus taeda) seed stratification, germination and post-germinative growth

Changes in the patterns of gene expression were examined during loblolly pine ( Pinus taeda L.) seed stratification, germination, and post-germinative growth. In both the megagametophyte and the embryo, DNA contents remained relatively constant at all stages examined. RNA contents, however, increased in both tissues following seed germination, particularly in the embryo where a 7-fold increase in the RNA content was observed 5 days after germination. Poly(A)⁺ RNA, extracted from megagametophytes and embryos, was translated in vitro in a rabbit reticulocyte lysate cell-free system. Analysis of [³⁵S]-methionine-labelled translation products by two-dimensional electrophoresis/fluorography indicated that there were changes in the populations of mRNAs during all developmental stages examined. In both the megagametophyte and the embryo several distinct mRNA populations, including one constitutively present at all stages examined, were identified. One mRNA population, present in the mature seed, decreased during seed stratification. Another population, not present in the mature seed, rose during the period of stratification that coincided with an increase in seed germinability. A third population, which appeared during seed germination, increased steadily during post-germinative growth. Besides these similarities, specific differences between megagametophyte and embryo were noted. For example, one mRNA population, which was present in the megagametophyte of the mature seed and remained constant during the stratification period, disappeared immediately following seed germination. In the embryo, one set of messages was germination specific. In total, these results show that mRNA populations change in a temporal fashion that is consistent with the patterns of de novo protein synthesis known to occur in loblolly pine during the same developmental periods.     

Seed‐based approach for identifying flora at risk from climate warming

In obligate seeding species, the germination niche is crucial for colonization and population survival. It is a high‐risk phase in a plant's life cycle, and is directly regulated by temperature. Seeds germinate over a range of temperatures within which there is an optimum temperature, with thresholds above and below which no germination occurs. We suggest that abrupt changes in temperature associated with a warming climate may cause a disconnect between temperatures seeds experience and temperatures over which germination is able to occur, rendering obligate seeding species vulnerable to decline and extinction. Using a bidirectional temperature gradient system, we examined the thermal constraints in the germination niche of some geographically restricted species from the low altitude mountains of the Stirling Range, southern Western Australia, including seedlots from lowland populations of four of these species. We demonstrated that high temperatures are not a limiting factor for germination in some restricted species, signifying a lack of relationship between geographic range size and breadth of the germination niche. In contrast, we identified other restricted species, in particular Sphenotoma drummondii, as being at risk of recruitment failure as a consequence of warming: seeds of this species showed a strong negative relationship between percentage germination and increasing temperature above a relatively low optimum constant temperature (13°C). We found some ecotypic differences in the temperature profiles between seeds collected from montane or lowland populations of Andersonia echinocephala, and while specific populations may become more restricted, they are perhaps at less risk of extinction from climate warming. This seed‐based approach for identifying extinction risk will contribute tangibly to efforts to predict plant responses to environmental change and will assist in prioritizing species for management actions, directing limited resources towards further investigations and can supplement bioclimatic modelling.     

Seed dispersal and germination patternsin a rare Mediterranean island endemic (Anchusa crispa Viv., Boraginaceae)

We analyse and discuss patterns of seed dispersal and germination of a rare endemic plant species, Anchusa crispa Viv. (Boraginaceae) from Corsica and Sardinia. This coastal plant inhabits dunes and back-dunes, and currently numbers only a few thousand individuals which generally occur in isolated populations. This study included experiments conducted in the field in Corsica and also under controlled conditions in the laboratory. Short-distance dispersal of A. crispa is performed by ants, both by myrmecochory and dyszoochory. The invasion of an exotic species of ant, Linepithema humile, could locally modify the dispersal system and possibly the population dynamics of A. crispa. Long-distance dispersal may occur by water transport since seeds can germinate after at least 1 week of immersion in seawater and readily float on the surface. Burial of seeds is favourable for germination, percentage germination being maximised at a depth of 1–2 cm below the soil surface. A. crispa has a seed bank of about ten viable seeds per m², which may contribute to the survival of this species which exists in small populations with a short life span. Due to its seed dispersal and germination patterns, the conservation of this species will necessitate that human disturbance, which can destabilise the surface of the sand is prevented and that new populations are introduced to favourable sites.     

Stable germination behavior but partly changing seed–seed interactions along a steep rainfall gradient

Germination marks a critical transition in plant life that is prone to high mortality. Strong selection pressure is therefore expected to finely tune it to environmental conditions. Our study on the common Mediterranean grass Brachypodium hybridum assessed whether germination behavior changes systematically along a steep natural rainfall gradient ranging from harsh desert to rather mild mesic-Mediterranean conditions. We specifically tested hypotheses that germination behavior confers greater risk-spreading in populations from drier, unpredictable environments, and that seeds from wetter populations are better competitors.In 14 populations (spanning 114–954 mm annual rainfall) we assessed three alternative key parameters of germination in a greenhouse experiment: between-year dormancy, days to emergence within a season, and temporal spread. Addition of neighbor seeds accounted for competition as another crucial environmental factor. In six of the 14 populations, we also compared seeds originating from corresponding north (more mesic) and south (more arid) exposed hill slopes to test whether germination patterns along the large-scale rainfall gradient are paralleled at this smaller scale.B. hybridum exhibited generally high germination fractions and rapid emergence with very little temporal spread, indicating overall little risk-spreading germination. Surprisingly, none of the three parameters changed systematically with increasing aridity, neither at large scale along the rainfall gradient nor at small scale between north and south exposures. Neighbor seeds, however, mildly suppressed germination. Germination of neighbor seeds, in turn, was more strongly suppressed by B. hybridum seeds from drier populations, and this effect was stronger for forb than for grass neighbor species.Our results provide strong evidence that increased risk-spreading germination is not a universal, essential strategy to persist in increasingly dry, unpredictable environments. They also highlight that competition with neighbors occurs even at the earliest plant life stage. Since neighbor effects were species-specific, competition among seeds can affect community composition at later plant stages.     

Management-driven evolution in a domesticated ecosystem

Millennia of human land-use have resulted in the widespread occurrence of what have been coined ‘domesticated ecosystems’. The anthropogenic imprints on diversity, composition, structure and functioning of such systems are well documented. However, evolutionary consequences of human activities in these ecosystems are enigmatic. Calluna vulgaris (L.) is a keystone species of coastal heathlands in northwest Europe, an ancient semi-natural landscape of considerable conservation interest. Like many species from naturally fire-prone ecosystems, Calluna shows smoke-adapted germination, but it is unclear whether this trait arose prior to the development of these semi-natural landscapes or is an evolutionary response to the anthropogenic fire regime. We show that smoke-induced germination in Calluna is found in populations from traditionally burnt coastal heathlands but is lacking in naturally occurring populations from other habitats with infrequent natural fires. Our study thus demonstrates evolutionary imprints of human land-use in semi-natural ecosystems. Evolutionary consequences of historic anthropogenic impacts on wildlife have been understudied, but understanding these consequences is necessary for informed conservation and ecosystem management.     

Variability on morphological and ecological seed traits of Limonium avei (De Not.) Brullo & Erben (Plumbaginaceae)

Limonium avei is an annual species occurring in the salt‐marshes and in limited surfaces of rocky areas around the Mediterranean coasts. Seed lots from five populations of this species, along a latitudinal gradient, were analyzed using an image analysis system to detect differences in seed morphology among populations. Germination requirements at constant (5–25°C) and alternating temperatures (25/10°C), both in light and in darkness, were evaluated for all populations, as well as the effect of the calyx removal on final seed germination and its rate. Morpho‐colorimetric analysis clearly identified seeds from different populations, habitats and substrates without misattributions among them. The calyx slowed the germination process, influencing both final germination and rate with respect to naked seeds. Seeds from all populations germinated with significantly higher percentages in the light, with respect to those incubated in the darkness, and showed rapid germination (time in days to reach 50% of germination: 0.5 days) at the warmer tested temperature (25°C). High germination (>80%) was also detected for seeds of all the investigated populations, except for those from the Apulian region (South Italy, ca. 60%). Our results highlight that L. avei has a high variability in seed morphology, probably habitat induced, and a fast germination response for all populations. Rapid germination may be an adaptive strategy that allows L. avei seeds to take advantage of transient favorable conditions during the germination stage, to ensure seedling establishment under the unpredictable rainfall pattern in the Mediterranean climate.     

The underground phase: a special challenge in studies of terrestrial orchid populations

Orchid seeds are minute and the first seedling stages of terrestrial species are underground, so there is little knowledge about the biology of the early life history and the size of seed/seedling populations relative to the number of plants that eventually emerge above ground. A recently developed field sowing technique makes it possible to explore a variety of topics such as the length of time that seeds spend as part of the soil seed bank, the phenology of seed germination, substrate requirements for germination, dependence of germination on fungal availability, substrate requirements and growth rate of seedlings, seedling phenology and development, and seedling-fungus specificity patterns in the soil. Several of these applications are relevant for conservation issues. Studies of Goodyera pubescens, Corallorhiza odontorhiza and Liparis lilifolia show considerable variation in germination strategies, patterns of seedling development and establishment, substrate requirements, and levels of specificity with the fungi.     

Response surfaces for the combined effects of heat shock and smoke on germination of 16 species forming soil seed banks in south‐east Australia

Abstract There is limited understanding of how fire‐related cues such as heat shock and smoke can combine to affect the germination response of seeds from fire‐prone vegetation because combinations of multiple levels of both cues have rarely been investigated. Germination response surfaces were determined for the combination of heat shock and smoke by applying factorial combinations of temperature (up to 100°C) and aerosol smoke (0–20 min) to 16 species that form soil seed banks in the Sydney region of south‐eastern Australia. Duplicate populations of three species were also examined to assess the constancy of a species response surface. Of the 19 populations examined, 16 showed a germination response to both the fire cues, which combined interactively in 14 populations, and independently in two. No population responded only to a single cue; however, seeds of 11 populations responded to heat in the absence of smoke, and nine responded to smoke in the absence of heat. Heat applied in the absence of smoke negatively affected germination in seven populations, either progressively as temperature increased, or above a set temperature. Negative germination responses over part of the temperature range were fully reversed at higher temperatures for unsmoked seeds of four populations (curvilinear heat response). Smoke effects were most frequently positive over all or part of the range of durations used, and when combined with heat frequently fully or partially reversed negative heat effects. Three populations required the obligatory combination of smoke and heat. A novel response to the cues was observed for three species, with smoke reversing negative heat effects at 75°C, being supplanted by a positive heat response of unsmoked seed at 100°C. The response surface for duplicate populations of two of the three species examined was variable. Heat shock and smoke frequently combined to affect germination, in both positive and negative ways. Consequently, to gain an accurate assessment of the response of seeds to fires, an experimental design that samples within the potential response zones of germination cues is essential.     

四川大头茶地理种群种子大小变异及对萌发、幼苗特征的影响

研究了四川大头萘６个分布地,７个地理种群种子大小变异特征、种子萌发及幼苗发育特征。结果表明,种子大小地理变异,南方种群种子变异较大,北方种上对较小,云南文山的种子最大最重,广西阳塑最轻最小,且最重约为最轻的２～３倍,不同种群种子萌发率也存在差异。南方种子差异较大,北方种子差异较小,大种子普遍有较高萌发率。种子太小小对苗生长速率、苗高、生物量（鲜重）有明显影响。     

Germination behaviour in populations ofCapsella bursa-pastoris (Cruciferae)

Germination behaviour of variousCapsella bursa-pastoris populations collected from Scandinavia, Middle Europe and the Alps, was tested in unheated, non-illuminated greenhouses (46 populations) and in growth chambers using 5–7 alternating temperature regimes (16 populations). For all populations, the influence of temperature on germination rate is straightforward: the higher the temperature, the greater the germination. Germination capacity, however, may depend on the geographical region. There is also a strong seed age effect on both, rate and capacity of germination. Once dormancy was broken, seeds from all populations were able to germinate over the entire range of temperatures. Some populations revealed a more or less pronounced temperature optimum for germination capacity, others germinated equally well over the entire temperature range. This indicates genetic heterogeneity between populations. However, no correlation between germinability and any environmental pattern was detected. The data indicate thatCapsella bursa-pastoris has adopted a germination strategy which includes a broad temperature tolerance. Germination of wildCapsella plants seems to be regulated by the factors contributing to the inception and breaking of dormancy which depend on pre- and postharvest conditions. Adaptation in germination behaviour inCapsella bursa-pastoris is different from that in other life history traits (flowering behaviour, growth form parameters).     

Influence of altitude on local adaptation in upland tree species from central Argentina

• Steep climatic gradients boost morphological and physiological adjustments in plants, with consequences on performance. The three principal woody species of the Sierras Grandes Mountains of central Argentina have marked differences in sapling performance along their altitudinal distribution. We hypothesize that the steep gradient of climatic conditions across the species’ altitudinal distribution promotes trait differences between populations of different altitudes that are inherited by the following generation.
• Seeds from different altitudes were exposed to three temperature regimes to assess differential germination responses. Saplings were then transplanted to a greenhouse to assess possible variations in attributes and performance after 18 months.
• The three species showed differences in germination responses to temperature among altitudes and/or in sapling attributes and performance. In Maytenus boaria and Escallonia cordobensis, germination success was higher under high temperatures for the highest‐altitude, whereas lower temperatures boosted germination of the lowest altitudes. Polylepis australis showed no differences in germination among temperature treatments. In the greenhouse, saplings of the three species from intermediate altitudes showed high performance, whereas the upper and lower populations seemed to be adjusted to tolerating more stressful conditions (i.e., lower temperatures at the upper end and water stress at the lower end), showing lower performance toward both altitudinal limits.
• These patterns agree with those described for saplings growing under field conditions, suggesting adjustments in response to environmental changes undergone by populations along the altitudinal range. The marked adjustments of populations to the local environment suggest a potentially high impact of climatic change on species distribution.

     

Seed weight and germination behavior of the submerged plant Potamogeton pectinatus in the arid zone of northwest China

Variation in seed weight is common within and among plant species, but few studies have attempted to document the pattern of seed weight and germination attributes for aquatic macrophytes at a large scale. This study examined within‐species variation in seed weight and germination attributes and the effects of environmental factors on seed traits of the submerged plant Potamogeton pectinatus in the arid zone of northwest China. Our results showed that the average seed weight was 0.24 g per 100 seeds with a coefficient of variation (CV) of 28.4% among the eight P. pectinatus populations. The total germination fraction of seeds of P. pectinatus was relatively poor, less than 35% in seven P. pectinatus populations, and the lowest germination percentage found was only 2%. There were significant differences in seed weight, time to onset of germination, and total germination fraction among the eight different populations. Hierarchical partitioning analysis showed a strongly positive correlation between seed weight and water temperature and pH. Seed weight and the maternal environmental factors significantly affected both time to initiation of germination and total germination fraction. Our results suggest that (1) seed weight variation in P. pectinatus primarily is the result of temperature variation during fruit development; (2) relatively poor germination fraction suggests that seeds are relatively unimportant in the short‐term survival of populations and that it may be another adaptive trait allowing plants to take place in the right place and at the right time, especially in harsh environment; and (3) variation in seed germination traits should be determined by local environmental and intrinsic factors that interact in a complex fashion.     

温度对高山植物紫花针茅种子萌发特性的影响

温度是影响种子萌发的重要因子,在气候变暖和草地退化的背景下,认识温度变化对紫花针茅（Stipa purpurea） 种子萌发的影响有重要意义。作者系统研究了6个恒温条件（5、10、15、20、25、30℃）和3个变温条件（5/15、10/20、15/25℃）对不同居群紫花针茅种子萌发特性的影响。结果表明,紫花针茅种子适宜的萌发温度范围为15~25℃,低于10℃或高于30℃的温度都会抑制种子的萌发;与对应的恒温条件相比,变温条件不能起到促进紫花针茅种子萌发的作用;不同居群紫花针茅种子萌发对温度的响应既有相同点,也有不同点;海拔高度与种子萌发率存在极显著负相关关系,种子萌发季土壤温度与种子萌发存在显著负相关关系。本研究将为合理的保护和恢复紫花针茅草地提供一定科学依据。