Populational dynamics of three annual species of alpine plants in the Rocky Mountains

Summary Survivorship, fecundity, and seed bank size were measured over two years in alpine populations of the annuals, Koenigia islandica, Polygonum confertiflorum, and P. douglasii. The major loss of individuals from all populations occurred between seed dispersal and germination. Survival of vegetative plants to maturity was high in all species, usually above 80%, and the average number of seeds produced per plant was less than 10 in all species in both years. Seed banks existed for all three species but were small since more than 80% of the viable seeds in the soil germinated in the spring. Both survival and fecundity were negatively correlated with density in P. confertiflorum. These patterns of population dynamics are similar to those found in many annuals of temperate environments.     

THE EFFECT OF SEED SIZE AND MATERNAL SOURCE ON INDIVIDUAL SIZE IN A POPULATION OF LUDWIGIA LEPTOCARPA (ONAGRACEAE)

Seed size is normally distributed for many annual species, while mature plant size is frequently positively skewed. A study was conducted to determine the influence of seed size and the role of genetic differences in determining relative seedling size for Ludwigia leptocarpa. Seed size had a significant effect on percentage germination and time of seed germination but no effect on dry weight or leaf area of seedlings. Seed size and spacing had a significant effect on seedling dry weight for plants grown under competition, while relative day of emergence had no effect. Familial (genetic) differences were found in average seed weight between maternal plants, but not in average number of days to germination, average weight of seeds which germinated, or shoot dry weight. It is concluded that neither seed size alone nor genetic differences between plants are directly responsible for the development of size hierarchies in Ludwigia leptocarpa populations. Large seed size does convey an advantage in growth when plants from seeds of differing initial size interact.     

A meta-analysis of the effects of frugivory (endozoochory) on seed germination: role of seed size and kind of dormancy

Heretofore, no study has determined how germination of ingested seeds is affected by the kind (class) of dormancy nor by seed dormancy x seed size interaction. Thus, we aimed to determine the effects of seed size, kind of dormancy and their interaction on germination of defecated seeds using a meta-analysis. We collected data for 366 plant species in 97 plant families from 76 publications. In general, gut passage significantly increased germination percentage of defecated seeds by 5% compared with that of control seeds. Germination percentages of non-dormant, physiologically dormant, and morphologically/morphophysiologically dormant seeds (all water-permeable) significantly decreased after gut passage by 40, 18, and 14%, respectively, compared with control seeds (non-gut-passed). Changes in germination percentage of seeds with physical dormancy (water-impermeable) were positive, and gut passage increased germination by 69% compared with control seeds. Germination of small seeds decreased 8% after gut passage, whereas germination of both medium and large seeds increased by 18%. However, changes in germination percentage differed between categories of seed size in each class of dormancy. In physically dormant seeds, germination of all seed sizes improved after gut passage, and the magnitude of increase was higher for large than for medium and small seeds. Thus, gut passage increased germination of medium-size water-permeable seeds (physiologically dormant and morphologically/morphophysiologically dormant) more than it did for large and small seeds. However, gut-passage decreased or did not change the germination percentage of non-dormant seeds. Seed size and kind of dormancy should be included in studies on the effect of gut passage on germination.     

Seed predation,pathogen infection and life-history traits in Brassica rapa

Herbivory and disease can shape the evolution of plant populations, but their joint effects are rarely investigated. Families of plants of Brassica rapa (Brassicaceae) were grown from seeds collected in two naturalized populations in an experimental garden. We examined leaf infection by the fungus Alternaria, seed predation by a gall midge (Cecidomyiidae) and plant life-history traits. Plants from one population had heavier seeds, were more likely to flower, had less fungal infection, had more seed predation and were more fecund. Fungal infection score and seed predation rate increased with plant size, but large plants still had the greatest number of undamaged fruits. Spatial heterogeneity in the experimental garden was significant; seed predation rate and fecundity varied among blocks. An apparent tradeoff existed between susceptibility to disease and seed predation: plants with the highest fungal infection score had the lowest seed predation rate. Alternaria infection varied between populations, but the disease had no effect on fecundity. Seed predation did reduce fecundity. Damaged fruits had 31.4% fewer intact seeds. However, evidence for additive genetic variation in resistance to seed predation was weak. Therefore, neither disease nor seed predation was likely to be a strong agent of genetically based fecundity selection.     

The role of the tolerance–fecundity trade‐off in maintaining intraspecific seed trait variation in a widespread dimorphic herb

Coexistence of species with different seed sizes is a long‐standing issue in community ecology, and a trade‐off between fecundity and stress tolerance has been proposed to explain co‐occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade‐off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade‐off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life‐cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance–fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size–number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.     

散枝猪毛菜的种子多型性及其萌发行为

散枝猪毛菜(Salsola bracchita)主要分布于新疆准噶尔荒漠,具有很强的抗干旱和抗盐碱能力。该文对散枝猪毛菜的果实进行了观察,表明散枝猪毛菜具有4种类型的散布单位和果实,这4种果实在形状、大小、颜色和着生方式上均有显著差异。A型果实绿色、球形、大,着生方式为横生,宿存花被革质,背部有紫红色翅状附属物,直径为(2.161±0.138) mm,单粒重为(3.810±0.113) mg;B型果实绿色、大、扁平、长圆形,着生方式为直立,宿存花被革质,背部有翅状附属物,长为(2.062±0.188) mm,宽为(1.720±0.148) mm,单粒重为(2.665±0.113) mg;C型果实绿色、大、扁平、长圆形,着生方式为直立,宿存花被膜质,背部无翅状附属物,长为(2.239±0.277) mm,宽为(1.844±0.150) mm,单粒重为(2.723±0.559) mg;D型果实黄色、小、扁平、长卵形,着生方式为直立,宿存花被膜质,背部无翅状附属物,长为(1.678±0.163) mm,宽为(1.390±0.110) mm,单粒重为(0.928±0.025) mg。A型、B型和C型种子(果实)在5 ℃/15 ℃、5 ℃/25 ℃、15 ℃/25 ℃(暗12 h/光12 h)变温条件下萌发率>68%,且B型和C型种子比A型种子有较高的萌发率和萌发速率。D型种子在3种变温条件下萌发缓慢,最终萌发率始终维持在较低水平(<20%),划破果皮和种皮能够显著提高D型种子的萌发率和萌发指数,表明D型种子处于生理休眠状态。散枝猪毛菜的种子多型性是对荒漠异质环境的一种适应。     

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.     

种子特征和播种深度对辽东栎种子萌发和幼苗生长的影响

该文在玻璃温室内的遮阴环境下,采用盆播方法研究了辽东栎种子特征(大小和种皮)和播种深度(0、3、6、10 cm)对种子萌发和幼苗生长的影响。结果表明:(1)辽东栎大种子萌发率、萌发速率、萌发值和萌发指数在所有播种深度均高于小种子。其中非去皮种子萌发率、萌发速率和萌发指数在0 cm播种深度的不同大小种子间差异显著,去皮种子所有萌发参数在6和10 cm播种深度的不同大小种子间均差异显著;去皮可促进大种子萌发,但抑制小种子萌发;不同大小种子所有萌发参数均在0 cm播种深度最大,在10cm播种深度最小。(2)不论有无种皮,大种子萌发幼苗的叶片数、单株叶面积、总干物质质量和根冠比在所有播种深度均大于小种子萌发幼苗;去皮种子萌发幼苗的株高、基径、叶片数、单株叶面积和总干物质质量在所有播种深度均不同程度地小于非去皮种子萌发幼苗,但前者根冠比在0、3、6 cm播种深度大于后者;随着播种深度增大,幼苗株高、叶片数、单株叶面积、总干物质质量和根冠比等生长参数均呈减小趋势,但基径随播种深度增大而增大。     

Seed yield,seed size and germination behaviour in the annualPogogyne abramsii

Summary Models of the evolution of seed dormancy reveal that dormancy is favoured either when opportunities for establishment vary over time and when there is wide variation in the probability of success, or when the probability of success is limited by frequency dependence. Empirical evidence supporting the temporal heterogeneity hypothesis exists, but there is scant evidence for dormancy being favoured by frequency dependent competition among seedlings. We test the hypothesis that the intensity of between-sib competition should favour a positive relationship between maternal fecundity and seed dormancy. This hypothesis is supported for the rare, vernal pool annual,Pogogyne abramsii: the proportion of dormant offspring was significantly higher among high fecundity mothers than among other mothers. Dormancy inP. abramsii is controlled by the seed coat, a maternal tissue, so delaying germination favours the inclusive fitness of mothers by reducing the potential for competition among siblings. Seed weight and time to first germination varied significantly amongP. abramsii plants and mean seed weight increased linearly with plant biomass. Seed weight and seed number are independently regulated by plant size. Overall, seed weight varied 10-fold and variability in seed weight within mothers was not explained by plant biomass, seed yield or mean seed weight. GerminableP. abramsii seeds were significantly heavier than dormant seeds, and germinable seeds heavier than 0.20 mg germinated more rapidly than those smaller than 0.20 mg.     

Pollen source affects female reproductive success and early offspring traits in the rare endemic plant Polemonium vanbruntiae (Polemoniaceae)

Understanding the relative magnitudes of inbreeding and outbreeding depression in rare plant populations is increasingly important for effective management strategies. There may be positive and negative effects of crossing individuals in fragmented populations. Conservation strategies may include introducing new genetic material into rare plant populations, which may be beneficial or detrimental based on whether hybrid offspring are of increased or decreased quality. Thus, it is important to determine the effects of pollen source on offspring fitness in rare plants. We established pollen crosses (i.e. geitonogamous‐self, autonomous‐self, intrasite‐outcross, intersite‐outcross and open‐pollinated controls) to determine the effects of pollen source on fitness (seeds/fruit and seed mass) and early offspring traits (probability of germination, number of leaves, leaf area and seedling height) in the rare plant Polemonium vanbruntiae. Open‐pollinated, intrasite‐outcross and geitonogamous‐self treatments did not differ in fitness. However, plants receiving autonomous‐self pollen had the lowest fitness and the lowest probability of seed germination. Intersite‐outcross plants contained fewer seeds/fruit, but seeds germinated at higher frequencies and seedlings were more vigorous. We also detected heterosis at the seed germination stage. These data may imply that natural populations of P. vanbruntiae exhibit low genetic variation and little gene flow. Evidence suggests that deleterious alleles were not responsible for reduced germination; rather environmental factors, dichogamy, herkogamy and/or lack of competition among pollen grains may have caused low germinability in selfed offspring. Although self‐pollination may provide some reproductive assurance in P. vanbruntiae, the result is a reduction in germination and size‐related early traits for selfed offspring.     

VARIATION IN SEED WEIGHT AND ITS EFFECTS ON GERMINATION IN PASTINACA SATIVA L. (UMBELLIFERAE)

Pastinaca sativa (wild parsnip) produces seeds on the primary, secondary, and tertiary umbels of the flowering stalk. Within plants, variation in seed weight is about twofold. Secondary and tertiary seed weight is 73% and 50% of primary seed weight, respectively. Maximum variation in seed weight between plants is sixfold when tertiary seeds from a small plant are compared to primary seeds from a large plant. Within an umbel order, variation in seed weight between plants is correlated with plant size. Under autumn germinating conditions in the laboratory, final germination of seeds from different umbel orders does not differ but smaller seeds germinate more rapidly than larger seeds. Under spring germination conditions in the laboratory, significantly more primary and secondary seeds germinate than tertiary seeds and the rate of germination is independent of seed weight. Field germination of seeds from different umbel orders produces similar results except that in the spring both secondary and tertiary seed germination is lower than that of primary seeds. These results suggest that with respect to seed germination characteristics small seeds may have a competitive advantage over large seeds in the autumn because they germinate more quickly, but in the spring small seeds are at a disadvantage because they have lower overall germination. Because most germination in the field occurs in the spring, population recruitment from small seeds is likely to be substanially less than that from large seeds.     

SEED MORTALITY IN DAUCUS CAROTA POPULATIONS: LATITUDINAL EFFECTS

Daucus carota, a common herbaceous weed, grows over a wide latitudinal range in eastern North America. Viability and germination tests of mature seeds collected from 36° to 45°N were conducted to measure predispersal seed mortality. Viability and germination declined as latitude of the seed source decreased. Only 30–50% of the seeds from southern populations germinated owing to high embryo inviability and absence of embryos. Sixty to ninety percent of the seeds from northern populations germinated. Reciprocal planting of seeds in outdoor experimental plots at three latitudes and testing of seeds over two generations together showed that the environment in which seeds mature, rather than environmental preconditioning over generations or genetically-based differences among populations, explain this variation in germination ability. Within-latitude germination declined in experimental plots as population age of the seed source within latitudes increased. The data indicate that predispersal seed mortality can influence local population persistence and that seed mortality is an increasingly important factor in population regulation at the southern limit of the species’ range.     

THE EFFECT OF SALINITY AND TEMPERATURE ON THE GERMINATION OF POLYMORPHIC SEEDS AND GROWTH OF ATRIPLEX TRIANGULARIS WILLD.

Polymorphic seeds of Atriplex triangularis were germinated at various temperatures (5–15 C, 5–25 C, 10–20 C, 20–30 C) and salinity regimes (0 to 1.5% NaCl) in order to determine their germinability and early seedling growth under these conditions. Larger seeds generally had a higher germination percentage in saline medium. The rate and percentage of germination decreased with increased salinity stress. A thermoperiod of 25 C day and 5 C night, 12 hr/12 hr, temperature enhanced germination of seeds. Early seedling growth is promoted in larger seeds at lower salinity, and at high-day and low-night temperatures. Polymorphic seeds have different physiological requirements which provide alternative situations for seed germination in natural habitats.     

Widespread seed limitation affects plant density but not population trajectory in the invasive plant Centaurea solstitialis

In some plant populations, the availability of seeds strongly regulates recruitment. However, a scarcity of germination microsites, granivory or density-dependent mortality can reduce the number of plants that germinate or survive to flower. The relative strengths of these controls are unknown for most plant populations and for exotic invaders in particular. We conducted a seed addition experiment with a granivore exclusion treatment in a field setting to explore how these factors interact to regulate populations of the widespread invader Centaurea solstitialis (yellow starthistle) at three study sites across the plant’s range in California. We coupled the experimental approach with observational studies within established C. solstitialis populations to estimate seed rain, recruitment and mortality at natural densities. Seed limitation occurred in both experimental and observational plots in all populations. Although vertebrate granivores were active at each site, they had no effect on C. solstitialis recruitment. Density increased mortality, but the effect was variable and weak relative to its effect on fecundity. The seed limitation that was evident at the seedling stage persisted to flowering. Seed-limited populations such as these ought to be highly sensitive to losses to seed predators, and many biological control agents, including those established for C. solstitialis, are seed predators. However, flowering plant density was decoupled from seed production by a strong compensatory response in the surviving plants; seed production was nearly constant in plots across all seed addition levels. Thus, flowering plant density is reduced by the established biocontrol agents, but seed production compensates to replace the population every generation, and no long-term decline is predicted.     

沙埋和种子大小对固沙禾草沙鞭的种子萌发与幼苗出土的影响

该文研究了野外条件下不同深度的沙埋对沙鞭(Psammochloa villosa)种子萌发和幼苗出土的影响,以及温室条件下种子大小对不同深度沙埋后的种子萌发和幼苗出土的影响。结果表明,沙埋深度显著影响沙鞭的种子萌发率、幼苗出土率和种子休眠率。沙子表面的种子不能萌发。2 cm的浅层沙埋时的种子萌发率和幼苗出土率最高,1 cm 沙埋的种子萌发率和幼苗出土率次之。沙埋深度超过2 cm之后,沙鞭的种子萌发率和幼苗出土率与沙埋深度呈负相关。2 cm的种子休眠率最低。从2 ~12 cm,种子休眠率随着沙埋深度的增加而增加。在幼苗能够出土的深度(1~6 cm),幼苗首次出土所需的时间随着沙埋深度的增加而延长。种子大小对沙鞭的种子萌发率没有显著影响。但是在深层沙埋(6 cm)时,与小种子相比,大种子产生的幼苗的出土率较高。从2~6 cm,大种子形成的幼苗的茎长度都较长。     

Herbivory and abiotic factors affect population dynamics of Arabidopsis thaliana in a sand dune area

Population dynamics of the annual plant Arabidopsis thaliana (L.) Heynh. were studied in a natural habitat of this species on the coastal dunes of the Netherlands. The main objective was to elucidate factors controlling population dynamics and the relative importance of factors affecting final population density. Permanent plots were established and plants were mapped to obtain data on survival and reproductive performance of each individual, with special attention to herbivore damage. In experimental plots we studied how watering, addition of nutrients, artificial disturbance, and natural herbivores affected survival and growth. Mortality was low during autumn and early winter and high at the time of stem elongation, between February and April. A key factor analysis showed a high correlation between mortality from February to April and total mortality. The specialist weevils Ceutorhyncus atomus and C. contractus (Curculionidae) were identified as the major insect herbivores on A. thaliana, reducing seed production by more than 40 %. These herbivores acted in a plant size-dependent manner, attacking a greater fraction of the fruits on large plants. While mortality rates were not affected by density, fecundity decreased with density, although the effect was small. Adding water reduced mortality in rosette and flowering plant stages. Soil disturbance did not increase seed germination, but did have a significant positive effect on survival of rosette and flowering plants. Seed production of A. thaliana populations varied greatly between years, leading to population fluctuations, with a small role for density-dependent fecundity and plant size-dependent herbivory.     

Germination ecology and seed population dynamics of Digitalis purpurea

Summary The germination ecology and the dynamics of the generative reproduction in populations of Digitalis purpurea L. were investigated in the field as well as in experiments. Germination of fresh seeds in the dark on moist filter paper appeared to differ between populations. These differences were eliminated when a moist natural soil functioned as germination substrate. An interaction between the spectral composition of light and the germination substrate was present. Germination in gradients of light, temperature and soil moisture revealed some clear-cut results. Germination proved to be strongly dependent on the percentage of vegetation cover. During two years of burial in litter bags, the number of buried viable seeds did not decrease. From one generation of seeds produced in a natural population, 18% was introduced into the buried seed bank, 10% germinated in autumn and 24% was present as a enforced dormant surface seed bank in late autumn.The results are discussed in relation to secondary succession. can be derived from Milton (1936), Salisbury (1942) and Thompson and Grime (1979). Soil disturbance and germination seem to be correlated in D. purpurea (Grime 1979). The purpose of this study is to analyse the dormancy and germination behaviour of D. purpurea in relation to the relevant environmental factors in order to explain the mechanisms of entry into, and the escape of D. purpurea seeds from a seed bank. Furthermore, an attempt will be made to quantify seed rain as well as the fate of different germinating and non-germinating seed rain fractions in space and time per unit area, in different stages of succession.     

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).     

Population fragmentation may reduce fertility to zero in Banksia goodii — a demonstration of the Allee effect

All individuals of all known populations of Banksia goodii were assessed for seed production. Small populations produced no or only a few seeds per unit canopy area. Effects of population size on seed production per unit area and seed production per plant were present over the whole range of population sizes, indicating that even in large populations seed production may still not be at its maximum. Resource differences could not explain this disproportionate decrease in seed production with decline in population size, because there were no differences in soil properties and understorey or overstorey cover between the small and large populations. Although plants in small and large populations were similar in size, seed production per plant was much lower in small populations. This was not because plants in small populations produced fewer cones but because the fraction of these cones that was fertile was much lower. Five of the nine smallest populations (<200 m²) produced no fertile cones over the last 10 years. The number of seeds per fertile cone did not depend on population size. The results are discussed in relation to pollination biology.     

Plant size,geitonogamy and seed set in Ipomopsis aggregata

Summary We used powdered fluorescent dyes to estimate receipt of self vs. outcross pollen in the self-incompatible species Ipomopsis aggregata (Polemoniaceae). Flowers on small and large plants received equal amounts of outcross pollen, whereas flowers on large plants received more self pollen, so the proportion of self pollen delivered through geitonogamy increased with plant size. In natural populations emasculation of all flowers on a plant raised average seed set per flower from 5.19 to 6.99 and also raised fruit set, though not significantly. From these results one expects a negative correlation between plant size and seeds per flower. The opposite trend was observed in a sample of plants in the field, suggesting that deleterious effects of geitonogamy on female fecundity in large plants can be overruled by other factors such as size-related fruit or seed abortion. Results are discussed in relation to the evolution of gynodioecy.