Do seed transfer zones for ecological restoration reflect the spatial genetic variation of the common grassland species Lathyrus pratensis?

A common ecological restoration approach is the reestablishment of vegetation using seed mixtures. To preserve the natural genetic pattern of plant species local seed material should be used. Consequently, seed transfer zones (seed production areas and seed provenance regions) have been delineated for ecological restoration in Germany. Although it is assumed that these transfer zones represent genetic variation, there remains a lack of empirical data. In this study, we analyzed whether seed transfer zones reflect the genetic variation of the common grassland species Lathyrus pratensis. We sampled 706 individuals from 37 populations in Bavaria, Germany and analyzed genetic variation using amplified fragment length polymorphisms. In our study, we observed higher levels of genetic variation and fragment rarity in the southern Bavarian populations compared to northern populations. Our analyses revealed a strong genetic differentiation between southern and northern Bavarian populations delineated along the Danube River. However, seed production areas and seed provenance regions reflected genetic variation of L. pratensis only to a limited degree. Our study illustrates that the level of genetic variation within populations strongly depends on population history. Furthermore, the geomorphological and climatic attributes, which have been used to delineate seed provenance regions, do not reduce gene flow among populations. Seed collections for gene banks and seed production should comprise seeds from populations in southern and northern Bavaria representing the strong genetic variation between both regions, but prioritize southern populations due to higher levels of variation.     

Responses of Primula vulgaris to light quality in the maternal and germination environments

• In the model species Arabidopsis thaliana phytochromes mediate dormancy and germination responses to seasonal cues experienced during seed maturation on the maternal plants. However, the effect of the maternal light environment on seed germination in native wild species has not been well studied. This is particularly important given its practical application in the context of environmental restoration, when there can be marked changes in the canopy.
• Plants of Primula vulgaris were grown in the field over two vegetative seasons under four shading treatments from low to high ratio of red to far‐red light (R:FR). Leaf and seed traits were assessed in response to the light treatments. The germination of seeds from these four maternal environments (pre‐dispersal) was investigated at seven light and five temperature treatments (post‐dispersal).
• Thinner leaves, larger leaf area and greater chlorophyll content were found in plants growing in reduced R:FR. Shading in the maternal environment led to increased seed size and yield, although the conditions experienced by the maternal plants had no effect on seed germination. Seeds responded strongly to the cues experienced in their immediate germination environment. Germination was always enhanced under higher R:FR conditions.
• The observed phenotypic trait variation plays a major role in the ability of P. vulgaris to grow in a wide range of light conditions. However, the increased germination capacity in response to a higher R:FR for all maternal environments suggests potential for seedling establishment under vegetative shade only in the presence of canopy gaps.

     

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.     

The role of Rhytidoponera metallica (Hymenoptera,Formicidae) in facilitating post‐fire seed germination of three ant‐dispersed legume species

The fire avoidance hypothesis proposes that a benefit of seed dispersal by ants (myrmecochory) is to protect seeds from being killed during fire and to facilitate post‐fire germination of seeds that require heat shock to break their physical dormancy. The aim of this study was to quantify the effect of fire and seed burial by a predominant seed‐dispersing ant, Rhytidoponera metallica (subfamily: Ectatomminae) on germination levels of three ant‐dispersed legume species (Pultenaea daphnoides, Acacia myrtifolia and Acacia pycnantha). Experimental burial of seeds within aluminium cans at a site prior to being burnt and at an adjacent unburnt site showed that fire increased germination levels, particularly for seeds buried at 1‐ and 2‐cm deep and that overall, germination levels differed among the three plant species. To quantify seed burial depths and post‐fire germination levels facilitated by R. metallica ants, seeds were fed to colonies prior to fire at the burnt and unburnt sites. Of the seeds buried within nests that were recovered, between 45% and 75% occurred within the upper 6 cm of the soil profile, although unexpectedly, greater percentages of seeds were recovered from the upper 0–2 cm of nests in the unburnt site compared with nests in the burnt site. Germination levels of buried seeds associated with R. metallica nests ranged from 21.2% to 29.5% in the burnt site compared with 3.1–14.8% in the unburnt site. While increased seed germination levels were associated with R. metallica nests following fire, most seeds were buried at depths below those where optimal temperatures for breaking seed dormancy occurred during the fire. We suggest that R. metallica ants may provide fire avoidance benefits to myrmecochorous seeds by burying them at a range of depths within a potential germination zone defined by intra‐ and inter‐fire variation in levels of soil heating.     

Fitness of reciprocal F1 hybrids between Rhinanthus minor and Rhinanthus major under controlled conditions and in the field

The performance of first‐generation hybrids determines to a large extent the long‐term outcome of hybridization in natural populations. F₁ hybrids can facilitate further gene flow between the two parental species, especially in animal‐pollinated flowering plants. We studied the performance of reciprocal F₁ hybrids between Rhinanthus minor and R. major, two hemiparasitic, annual, self‐compatible plant species, from seed germination to seed production under controlled conditions and in the field. We sowed seeds with known ancestry outdoors before winter and followed the complete life cycle until plant death in July the following season. Germination under laboratory conditions was much lower for the F₁ hybrid formed on R. major compared with the reciprocal hybrid formed on R. minor, and this confirmed previous results from similar experiments. However, this difference was not found under field conditions, which seems to indicate that the experimental conditions used for germination in the laboratory are not representative for the germination behaviour of the hybrids under more natural conditions. The earlier interpretation that F₁ hybrid seeds formed on R. major face intrinsic genetic incompatibilities therefore appears to be incorrect. Both F₁ hybrids performed at least as well as and sometimes better than R. minor, which had a higher fitness than R. major in one of the two years in the greenhouse and in the field transplant experiment. The high fitness of the F₁ hybrids confirms findings from naturally mixed populations, where F₁ hybrids appear in the first year after the two species meet, which leads to extensive advanced‐hybrid formation and introgression in subsequent generations.     

植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。     

Seed‐enhancement combinations improve germination and handling in two dominant native grass species

The use of native grasses for both restoration and commercial purposes is becoming increasingly important globally. Many native grasses have limited success in seed‐based restoration (e.g. post‐mine rehabilitation) and commercial industries (e.g. agriculture) due to poor seed germination and handling. Seed‐enhancement technologies can assist in overcoming these barriers. This study aimed to use combinations of seed enhancements to overcome the germination and handling challenges in two dominant Australian native grass species with demand in restoration and commercial industries (Triodia wiseana C.A Gardner and Rytidosperma caespitosum [Gaudich.] Connor & Edgar). Selected enhancements included hydropriming (including inoculation with karrikinolide [KAR₁]), flash flaming, and seed coating. Combinations of these seed enhancements allowed improvements in both germination (by up to 55% and 18% for T. wiseana and R. caespitosum, respectively) and floret geometry (flowability, as measured through a mechanized seeder, improved by up to 6‐fold and 17‐fold for T. wiseana and R. caespitosum, respectively), with the order of enhancement application being important. The responses of each species to enhancements corresponded with key biological processes and ecological cues required for recruitment events in nature, such as fire and rainfall events. Triodia wiseana germination was driven by fire‐related cues (i.e. KAR₁, flaming), while R. caespitosum germination was highest in response to moisture‐related cues (i.e. hydro‐priming). Responses to seed enhancements (and combinations of) may have implications for the management and scaled use of the targeted species. This can assist in improving the restoration and commercial success of the study species, and potentially other grasses with germination and handling challenges, into the future.     

Effect of variation in self-incompatibility on pollen limitation and inbreeding depression in Flourensia cernua (Asteraceae) scrubs of contrasting density

Background and Aims

Selection may favour a partial or complete loss of self-incompatibility (SI) if it increases the reproductive output of individuals in the presence of low mate availability. The reproductive output of individuals varying in their strength of SI may also be affected by population density via its affect on the spatial structuring and number of S-alleles in populations. Modifiers increasing levels of self-compatibility can be selected when self-compatible individuals receive reproductive compensation by, for example, increasing seed set and/or when they become associated with high fitness genotypes.

Methods

The effect of variation in the strength of SI and scrub density (low versus high) on seed set, seed germination and inbreeding depression in seed germination (δ_germ) was investigated in the partially self-incompatible species Flourensia cernua by analysing data from self-, cross- and open-pollinated florets.

Key Results

Examination of 100 plants in both high and low scrub densities revealed that 51% of plants were strongly self-incompatible and 49 % varied from being self-incompatible to self-compatible. Seed set after hand cross-pollination was higher than after open-pollination for self-incompatible, partially self-incompatible and self-compatible plants but was uniformly low for strongly self-incompatible plants. Strongly self-incompatible and self-incompatible plants exhibited lower seed set, seed germination and multiplicative female fitness (floral display × seed set × seed germination) in open-pollinated florets compared with partially self-incompatible and self-compatible plants. Scrub density also had an effect on seed set and inbreeding depression: in low-density scrubs seed set was higher after open-pollination and δ_germ was lower.

Conclusions

These data suggest that (a) plants suffered outcross pollen limitation, (b) female fitness in partially self-incompatible and self-compatible plants is enhanced by increased mate-compatibility and (c) plants in low-density scrubs received higher quality pollen via open-pollination than plants in high-density scrubs.Key words: Flourensia cernua, population density, seed set, seed germination, female fitness, partial self-incompatibility, Mapimí Biosphere Reserve     

Hybrid zones between invasive Rorippa austriaca and native R. sylvestris (Brassicaceae) in Germany: ploidy levels and patterns of fitness in the field

Hybrid zones may serve as natural laboratories for evolutionary studies. One common viewpoint is that hybrids may always be less fit than their parents due to genetic discontinuities. An alternative idea is that genotype-environment interactions influence the outcome of natural hybridization. Our comparative study of two different natural hybrid zones between the invasive diploid Rorippa austriaca and the native polyploid R. sylvestris in Germany identified the ploidy level as a major determinant of hybrid fitness. Different ploidy levels and patterns of fitness were detected in different hybrid zones. In one hybrid zone (Mülheim, Ruhr valley) hybrids were pentaploid and showed a relatively high seed set, whereas in the second hybrid zone (Randersacker, Main valley) hybrids were triploid and displayed extremely low fitness values. Analyses of fitness values in different natural hybrid zones between the same two species may lead to very different conclusions about the evolutionary significance of natural hybridization.     

Wait or escape? Contrasting submergence tolerance strategies of Rorippa amphibia,Rorippa sylvestris and their hybrid

Background and Aims

Differential responses of closely related species to submergence can provide insight into the evolution and mechanisms of submergence tolerance. Several traits of two wetland species from habitats with contrasting flooding regimes, Rorippa amphibia and Rorippa sylvestris, as well as F₁ hybrid Rorippa × anceps were analysed to unravel mechanisms underlying submergence tolerance.

Methods

In the first submergence experiment (lasting 20 d) we analysed biomass, stem elongation and carbohydrate content. In the second submergence experiment (lasting 3 months) we analysed survival and the effect of re-establishment of air contact on biomass and carbohydrate content. In a separate experiment we analysed expression of two carbohydrate catabolism genes, ADH1 and SUS1, upon re-establishment of air contact following submergence.

Key Results

All plants had low mortality even after 3 months of submergence. Rorippa sylvestris was characterized by 100 % survival and higher carbohydrate levels coupled with lower ADH1 gene expression as well as reduced growth compared with R. amphibia. Rorippa amphibia and the hybrid elongated their stems but this did not pay-off in higher survival when plants remained submerged. Only R. amphibia and the hybrid benefited in terms of increased biomass and carbohydrate accumulation upon re-establishing air contact.

Conclusions

Results demonstrate contrasting ‘escape’ and ‘quiescence’ strategies between Rorippa species. Being a close relative of arabidopsis, Rorippa is an excellent model for future studies on the molecular mechanism(s) controlling these strategies.     

Seed set in Sorghum stipoideum,and not fire,determines the timing of breeding by Gouldian finches (Erythrura gouldiae)

Seeds of native grasses are an important food source for granivorous finches throughout the tropical savannas of northern Australia. The iconic Gouldian finch (Erythrura gouldiae), a threatened species endemic to these savanna grasslands, relies almost exclusively on the grass seeds of annual Sorghum spp. when breeding, and appears to time breeding with the availability of these seeds. Fire is common throughout the savanna grasslands of northern Australia and has the potential to alter Sorghum spp. seed germination and plant reproductive phenology which in turn could alter the Gouldian finch breeding phenology window. This study examines if the temporal relationship between breeding by Gouldian finches in the north‐east Kimberley region of Australia relates to fire and Sorghum stipoideum seed phenology. The availability of S. stipoideum seed was monitored at Gouldian finch breeding sites in association with local fire history and Gouldian finch breeding data. The effect of experimental fire (heat and smoke) on S. stipoideum seed dormancy and germination was also investigated. We found that the first heavy rainfalls preceded germination of S. stipoideum in November/December. Synchronous seed set by S. stipoideum plants the following March/April coincided with the start and duration of the Gouldian finch breeding season. The timing of dry season fires had no relationship with seed dormancy or the phenology of seed germination and seed set. Nor did the effects of smoke or heat affect seed dormancy and germination. These findings support the importance of the seed phenology of annual grass species, such as S. stipoideum, to breeding Gouldian finches, and also suggest that the occurrence of fire at a breeding site in the previous year does not alter the S. stipoideum seed or finch breeding window. The implications of these results for threatened Gouldian finch ecology and management are discussed in relation to previously published fire impacts on S. stipoideum seed nutrition and finch breeding site selection.     

Seed quality in hybrid swarm populations of Pinus mugo Turra and P. sylvestris L.

Crop characteristics involving cone size, average number of seeds per cone, and seed germinability have been studied for three hybrid swarm populations of Pinus mugo and P. sylvestris, two pure P. mugo populations and one pure P. sylvestris population, in northern Slovakia. Generally, reproduction capability characteristics were significantly reduced in hybrid swarms in relation to the control populations. Although there were no consistent differences between hybrid swarms and control populations in the average number of seeds per cone or cone size, seed germination rates and full seed production in hybrid swarms were lower than for the control population of P. sylvestris and one of the pure populations of P. mugo. Based on these data a conclusion has been drawn postulating partial rather than full fertility of the hybrid swarm populations of P. mugo and P. sylvestris.     

Cryptic seed heteromorphism in Packera tomentosa (Asteraceae): differences in mass and germination

Seed mass variation and heteromorphism may afford plant species differential germination behavior and ultimately seedling success, particularly in disturbed habitats. We asked whether such variation occurs in Packera tomentosa (Michx.) C. Jeffrey (Asteraceae), a clonal species of the southeastern USA. Seed mass was compared within and among genetic individuals differentiated using amplified fragment length polymorphisms. We compared central and peripheral seeds produced by disc and ray florets, respectively, for their morphology, mass, and germination behavior, including response to water availability, aging, and cold stratification. Seed mass was highly variable both within and among individuals and influenced germination behavior. We found cryptic seed heteromorphism in P. tomentosa. Central and peripheral seeds had similar morphologies but dissimilar mass and biomass allocation. We used failure time analysis to detect different germination behavior. Central seeds were heavier, contained larger embryos, and germinated faster and at a higher proportion in most germination studies. Highly variable mass and heteromorphism of seeds may allow persistence of P. tomentosa in its disturbed habitats. Based on our results, some future studies of Asteraceae species with disc and ray florets may need to account for possible differences between seed types, even when morphological differences are not apparent. Evaluation of individual seed mass and maternal differences in germination studies may assist in the detection of cryptic seed heteromorphism, a phenomenon thought to be common, yet rarely documented.     

Optimising seed processing techniques to improve germination and sowability of native grasses for ecological restoration

• Grasslands across the globe are undergoing expansive degradation due to human impacts and climate change. If restoration of degraded native grassland is to be achieved at the scale now required, cost‐effective means for seed‐based establishment of grass species is crucial. However, grass seeds present numerous challenges associated with handling and germination performance that must be overcome to improve the efficiency of seeding. Previous research has demonstrated that complete removal of the palea and lemma (husk) maximises germination performance, hence we investigated the effects of complete husk removal on seed handling and germination of four temperate Australian grass species.
• Three techniques were tested to remove the husk – manual cleaning, flaming or acid digestion (the latter two followed by a manual cleaning step); these techniques were refined and adapted to the selected species, and germination responses were compared.
• The complete removal of the husk improved seed handling and sowability for all species. Germination was improved in Microlaena stipoides by 19% and in Rytidosperma geniculatum by 11%. Of the husk removal methods tested, flaming was detrimental to seed germination and fatal for one species (R. geniculatum). Compared to manual cleaning, sulphuric acid improved the overall efficacy of the cleaning procedure and increased germination speed (T50) in Austrostipa scabra, Chloris truncata and M. stipoides, and improved final germination in R. geniculatum by 13%.
• The seed processing methods developed and tested in the present study can be applied to grass species that present similar handling and germination performance impediments. These and other technological developments (seed coating and precision sowing) will facilitate more efficient grassland restoration at large scale.

     

Bottleneck-induced dissolution of self-incompatibility and breeding system consequences in Aster furcatus (Asteraceae)

Aster furcatus is a rare species with extremely limited genetic variation at isozyme loci. We utilized crossing experiments and seed set data obtained from natural populations to verify that there is also little allelic variation at the self-incompatibility (S) locus. Seed set in several populations was limited by a low number of S-alleles. Associated with a low number of S-alleles in populations was the dissolution of the incompatibility system, manifest by individual variation in self-compatibility, and by complex dominance relationships among S-alleles. Plant self-compatibility was correlated with mean number of ovules per inflorescence. Thus, self-compatibility appeared to be under partial environmental influence. Computer simulations revealed that the shapes of seed set distribution curves of modeled self-incompatible plant populations depend on the number of incompatibility alleles in the populations. By varying the number of S-alleles in modeled populations, we generated seed set distribution curves similar to those of natural populations. Genetic bottlenecks reduce the number of S-alleles and the proportion of compatible matings in populations of multiallelic self-incompatible species. Self-compatible genotypes are at a selective advantage in populations that lack a sufficient number of S-alleles to produce compatible crosses. Aster furcatus appears to be evolving self-compatibility as a result of bottleneck-induced losses of S-alleles.     

Heteromorphic seeds of wheat wild relatives show germination niche differentiation

• Crop wild relatives are fundamental genetic resources for crop improvement. Wheat wild relatives often produce heteromorphic seeds that differ in morphological and physiological traits. Several Aegilops and Triticum species possess, within the same spikelet, a dimorphic seed pair, with one seed being larger than the other. A comprehensive analysis is needed to understand which traits are involved in seed dimorphism and if these aspects of variation in dimorphic pairs are functionally related.
• To this end, dispersal units of Triticum urartu and five Aegilops species were X‐rayed and the different seed morphs weighed. Germination tests were carried out on seeds, both dehulled and left in their dispersal units. Controlled ageing tests were performed to detect differences in seed longevity among seed morphs, and the antioxidant profile was assessed in terms of antioxidant compounds equipment and expression of selected antioxidant genes. We used PCA to group seed morphs sharing similar patterns of germination traits, longevity estimates and antioxidant profile.
• Different seed morphs differed significantly in terms of mass, final germination, germination timing, longevity estimates and antioxidant profile in most of the tested species. Small seeds germinated slower, had lower germination when left in their dispersal units, a higher antioxidant potential and were longer‐lived than large seeds. The antioxidant gene expression varied between morphs, with different patterns across species but not clearly reflecting the phenotypic observations.
• The results highlight different trait trade‐offs in dimorphic seeds of Aegilops and T. urartu, affecting their germination phenology and longevity, thereby resulting in recruitment niche differentiation.

     

Post‐Dispersal Seed Removal and Germination Selected Tree Species Dispersed by Cebus capucinus on Barro Colorado Island,Panama1

Dispersal quality, an important component of seed disperser effectiveness, may strongly affect the rate of plant recruitment. Here we evaluated the quality of Cebus monkey dispersal by comparing the secondary removal fate and germination of fresh and Cebus‐ingested seeds of nine tree species on Barro Colorado Island, Panama. Overall, rates of secondary seed removal by vertebrates were low, with most Cebus defecations remaining undisturbed for extended periods on the forest floor. Only four of 30 feces were completely buried by dung beetles, and we found significantly higher vertebrate removal of defecated seeds than control seeds for only one species, Cordia bicolor. Seed germination varied greatly between plant taxa. Seeds of 3 out of 9 species showed significantly higher percent germination after monkey gut passage than control fresh seeds. Germination times tended to be shorter for defecated than for control seeds but were significantly different only for one of nine species, Cecropia insignis. Low rates of seed removal from Cebus feces, coupled with high germination probabilities, suggest high dispersal effectiveness for Cebus and contrasts strongly with patterns of post‐dispersal seed fate recorded for other primate species.     

Outcrossing rates in two self‐compatible,hybridising Rhinanthus species: implications for hybrid formation

The congeners Rhinanthus angustifolius and Rhinanthus minor, two annual hemiparasites pollinated by bumblebees, are known to hybridise in the wild. Both species are self‐compatible, but the capacity for autonomous selfing is higher in R. minor. This suggests a difference in realized outcrossing rates, which have not been determined before in these species. Using microsatellites, both species turned out to have mixed mating systems, but with a much lower multilocus outcrossing rate (0.13) for R. minor compared to R. angustifolius (0.76). We hypothesised that a higher outcrossing rate should lead to a higher chance of heterospecific pollination, and we therefore determined the rate of hybrid formation on each species in an artificial mixed population. Hybrid seeds were produced at low frequency (4.5%), and no significant difference was found between the species. It is therefore likely that post‐pollination processes influence hybrid seed formation to counteract the expected difference in heterospecific pollen deposition. We checked fruit set, seed set and the rate of autonomous selfing in controlled crosses in the greenhouse in 2 years, and found that fruit set (2003) or seed set (2010) were lower in R. angustifolius × R. minor crosses relative to the reciprocal cross. Hybrid seeds produced on R. angustifolius also had a much lower germination rate, so most of the established F₁ hybrid plants have the R. minor cytoplasm. The formation of advanced hybrids depends on pollinator preference, which is biased towards R. angustifolius if present in sufficient numbers, because it offers more rewards.     

Seed size and seed germination in the Mediterranean fire-prone shrub <Emphasis Type="Italic">Cistus </Emphasis><Emphasis Type="Italic">ladanifer</Emphasis>

The germination response of different sized seeds from individuals of a Mediterranean fire-prone shrub (Cistus ladanifer) was investigated in relation to pre-germination heating. A control (no heating), a low temperature during a short exposure time (50°C during 5 min), a high temperature during a short exposure time (100°C during 5 min) and a high temperature during a long exposure time (100°C during 15 min) were applied to seeds from different individual plants with different mean seed weight. These pre-germination treatments resemble natural germination scenarios for the studied species, absence of fire, low intensity pasture fire, typical Mediterranean shrub fire, and severe fire with high fuel load. Mean seed weight only showed a marginally significant positive correlation with the proportion of germinated seeds whatever the pre-germination treatment. These results suggest that seed dormancy is unrelated to seed size and that under the experimental conditions used in this study, the effect of seed size on seed germination is low. Nevertheless, larger seeds could be favoured in natural conditions, especially under the high competition scenario which arise after wildfires. Control seeds showed a negative correlation between seed size and germination velocity suggesting that lighter seeds could take advantage from early germination in recruitment events in the absence of wildfires. Nevertheless, even the lower pre-germination heating treatment turns this correlation in not significant, suggesting a strong selection pressure (unrelated to seed size) for early germination after fire events. In our study, different sized seeds of C. ladanifer seem to perform better under different germination scenarios suggesting that seed size variation could be maintained by the alternation of recruitments without wildfires and recruitments after wildfire events.     

GENETIC EFFECTS OF GERMINATION TIMING AND ENVIRONMENT: AN EXPERIMENTAL INVESTIGATION

Seeds of many species do not germinate immediately after dispersal, but instead may remain indefinitely in a dormant but viable state. Although it is well established that seeds often exhibit diversified patterns of dormancy and germination, the causes and consequences of this variation remain poorly understood. In this study, we investigate the extent to which seed genotypes of the desert mustard Lesquerella fendleri differentially germinate and establish under experimental conditions in a greenhouse. We used a two-way factorial design to compare genotypes of Lesquerella plants derived from seeds that germinated and established at different times and under different soil water regimes. Overall allozyme allele frequencies of Lesquerella plants varied significantly with both germination time and initial soil water availability. Single-locus heterozygosity analyses revealed that seeds sown into initially low water conditions produced plants that were significantly more heterozygous than plants derived from seeds experiencing constantly high water conditions, but heterozygosity did not differ significantly among plants originating from early- and late-germinating seeds. This is the first study to experimentally demonstrate that germination timing and environment can significantly affect the genetic structure of emerging plant populations. The study suggests that germination and survival behavior may (1) play an important role in generating and maintaining the genetic structure of natural plant populations and (2) set the stage for subsequent evolution.