Allozyme variation in allotetraploid Saxifraga osloensis and its diploid progenitors

The Scandinavian endemic plant Saxifraga osloensis is believed to be an allotetraploid derived from the diploids S. adscendens and S. tridactylites. We used gel electrophoresis of enzymes to characterize the genetic constitution of the three species. The results support both the allotetraploid nature of S. osloensis and its ancestry from the two assumed parental species. The pattern of genetic differences among local populations of the three species indicates that S. osloensis has arisen from hybridization in the western part of its present distribution area, possibly at a single occasion.     

Seed germination and plant fitness response of a narrowly endemic,rare winter annual to spatial heterogeneity in microenvironment

Environmental heterogeneity affects distributions of plant species, although the effects of fine-scale heterogeneity on plant demographies are not widely studied. Diminutive winter annuals, especially rare taxa, can be sensitive to spatial variation in microenvironment as a consequence of their small stature above and belowground. To address whether spatial environmental heterogeneity affects demography, germination and fitness of Chorizanthe orcuttiana, an endangered winter annual distributed in distinct easterly and westerly microhabitats within an exceedingly narrow niche in California, we performed multiyear observational and empirical studies. We manipulated after-ripening environment, soil moisture and ambient light at both aspects, and profiled microclimate, soil physicochemistry and soil microbiomes at all sites. We show that easterly aspects host larger plants in larger populations, and have lower air temperatures combined with higher soil moisture in comparison to the west-facing sites. Yet, soil physicochemistry and microbiomes were similar across all sites. Manipulations of after-ripening conditions showed that seeds exposed to low humidity (17%) during dormancy and sown at easterly aspects exhibited the highest germination percentages, whereas seeds incubated in situ and subsequently sown at westerly aspects yielded the lowest germination. Simultaneous manipulations of soil moisture and light showed that at both aspects higher moisture combined with shade led to higher germination, whereas ambient soil moisture combined with shade yielded the lowest germination. Altogether, our studies show that the diminutive, rare winter annual C. orcuttiana exhibits higher germination and plant fitness under cooler soil conditions with higher soil moisture while preferring drier environments during after-ripening.     

Clonality and recombination in the arctic plant Saxifraga cernua

The circumarctic clonal plant Saxifraga cernua reproduces efficiently via bulbils, largely depends on insects for pollination and appears to set seed very rarely. However, high levels of genotypic variation observed at small spatial scales in the arctic archipelago of Svalbard have been taken as evidence of occasional sexual reproduction. Here we assess the relative contributions of mutation and recombination to random amplified polymorphic DNA variation in four populations in East Greenland and re-analyse the Svalbard data. Greater variation due to recombination was predicted in Greenland than in Svalbard, because the higher summer temperatures and longer growing season likely increase the chances for sexual reproduction. Although we observed higher levels of genotypic diversity in Greenland than in Svalbard, matrix incompatibility and linkage disequilibrium measures provided no evidence of more sexuality, suggesting differences in glacial/postglacial history. The genetic structure and spatial distribution of clones suggest that clonal migration may increase variability in local populations, which is consistent with frequent large-scale migration in this species inferred from a circumarctic analysis of chloroplast DNA haplotypes. We conclude that a combination of somatic mutations and sexual reproduction has contributed to the observed patterns of genotypic diversity in the Greenland and Svalbard populations of S. cernua , and that sexual reproduction is important in maintaining genotypic diversity, despite the rarity of observations of seed setting.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 209–217.     

Bet hedging in desert winter annual plants: optimal germination strategies in a variable environment

In bet hedging, organisms sacrifice short‐term success to reduce the long‐term variance in success. Delayed germination is the classic example of bet hedging, in which a fraction of seeds remain dormant as a hedge against the risk of complete reproductive failure. Here, we investigate the adaptive nature of delayed germination as a bet hedging strategy using long‐term demographic data on Sonoran Desert winter annual plants. Using stochastic population models, we estimate fitness as a function of delayed germination and identify evolutionarily stable strategies for 12 abundant species in the community. Results indicate that delayed germination meets the criteria as a bet hedging strategy for all species. Density‐dependent models, but not density‐independent ones, predicted optimal germination strategies that correspond remarkably well with observed patterns. By incorporating naturally occurring variation in seed and seedling dynamics, our results present a rigorous test of bet hedging theory within the relevant environmental context.     

Hydrothermal thresholds for seed germination in winter annual forbs from old‐field Mediterranean landscapes

• Under Mediterranean climates with dry‐hot summers and cool‐wet winters, many forbs with potential for habitat restoration are winter annuals, but there is little information about their germination.
• We performed laboratory germination experiments on 13 ruderal dicots native to Andalusia (southern Spain). We measured the germination of recently harvested seeds from natural populations across nine temperature treatments (from 5 to 35 °C, constant and alternate); two storage periods; and eight water stress treatments (from 0 to ?1.0 MPa). We then calculated the hydrothermal thresholds for seed germination.
• Final germination ranged from 0–100% and results were mixed in response to temperature. Base temperature was below 6 °C, optimal temperature was around 14 °C and the ceiling temperature around 23 °C. For five species, 10 months of storage improved total germination, indicating a dormancy‐breaking effect, but the other species did not respond or had their germination reduced. All species were relatively tolerant to water stress, with base water potential ranging from ?0.8 to ?1.8 MPa.
• Our results suggest that hydrothermal germination thresholds, rather than physiological dormancy, are the main drivers of germination phenology in annual forbs from Mediterranean semi‐dry environments. The variation in germination responses of these forb species differs from winter annual grasses, but their seeds are all suitable for being stored before restoration.

     

Chemotaxonomic importance of diarylheptanoids and phenylpropanoids in Saxifraga tangutica (Saxifragaceae)

A phytochemical investigation of Saxifraga tangutica led to the isolation of 11 compounds, including eight diarylheptanoids (1–6, 10 and 11) and three phenylpropanoids (7–9). The chemical structures were established by extensive analysis of their MS and NMR spectroscopic data or comparison with literature data. In the present research, we report the isolated compounds 1–11, for the first time, in the species S. tangutica. Moreover, compounds 1, 2 and 4–11 have not been reported from any species in Saxifragaceae family. Furthermore, we discuss the chemotaxonomic significance of the isolated compounds.     

Genetic enrichment of the arctic clonal plant Saxifraga cernua at its southern periphery via the alpine sexual Saxifraga sibirica

Isolation of populations at the margins of a species range may lead to decreasing genetic diversity via genetic drift and inbreeding. Hybridization between peripheral populations of two species can, however, counteract genetic impoverishment. The mainly clonal, polyploid plant Saxifraga cernua has a wide arctic distribution but also extends southwards into alpine sites. In the Ural Mountains, its peripheral distribution overlaps with that of its sexually reproducing, diploid relative Saxifraga sibirica, and fertile polyploids of more or less intermediate appearance are found in this overlap zone. We used amplified fragment length polymorphism (AFLP) analysis to address the potential impact of interspecific gene flow on genetic diversity in the peripheral populations. A total of 149 plants from 17 populations along a 1650 km south-north gradient were analysed for 253 markers. The results suggest that three Middle Ural populations containing fertile and morphologically more or less intermediate plants have been affected by hybridization. All of these plants formed a strongly supported (100%) group with S. cernua in a neighbour-joining tree, but their AFLP phenotypes assigned either to S. cernua or to artificial (simulated) F(1) hybrids between S. cernua and S. sibirica in multilocus assignment tests. The three populations were highly diverse with virtually every plant representing a distinct AFLP phenotype, providing additional evidence for formation of later-generation hybrids and/or backcrossing to S. cernua. In contrast, other peripheral populations of S. cernua were typically monoclonal, suggesting that hybridization with S. sibirica can increase genetic diversity in S. cernua at its southern periphery.     

Variable demography in relation to germination time in the annual plant Tagetes micrantha Cav. (Asteraceae)

This study tests whether variability in a life history trait such as emergence time has ecological and evolutionary significance in seasonal environments due to their relationship with fitness components. The population dynamics of nine different cohorts of seedlings emergence were analyzed in a natural population of the annual herb Tagetes micrantha Cav. (Asteraceae). Temporal variation in seedling emergence was related to the pattern of precipitation, while spatial variation could be related to topographical heterogeneity. Plants that emerged at the beginning of the season (early germination) had lower probabilities of survivorship than those that emerged at the end of the season (late germination). In contrast, plants that emerged early had, on average, higher fecundity than those that emerged late. The net result of these contrasting patterns of survival and fecundity constitute a life history trade-off in T. micrantha.     

The relative importance of local conditions and regional processes in structuring aquatic plant communities

1. The structure of biological communities reflects the influence of both local environmental conditions and processes such as dispersal that create patterns in species’ distribution across a region. 2. We extend explicit tests of the relative importance of local environmental conditions and regional spatial processes to aquatic plants, a group traditionally thought to be little limited by dispersal. We used partial canonical correspondence analysis and partial Mantel tests to analyse data from 98 lakes and ponds across Connecticut (northeastern United States). 3. We found that aquatic plant community structure reflects the influence of local conditions (pH, conductivity, water clarity, lake area, maximum depth) as well as regional processes. 4. Only 27% of variation in a presence/absence matrix was explained by environmental conditions and spatial processes such as dispersal. Of the total explained, 45% was related to environmental conditions and 40% to spatial processes. 5. Jaccard similarity declined with Euclidean distance between lakes, even after accounting for the increasing difference in environmental conditions, suggesting that dispersal limitation may influence community composition in the region. 6. The distribution of distances among lakes where species occurred was associated with dispersal‐related functional traits, providing additional evidence that dispersal ability varies among species in ways that affect community composition. 7. Although environmental and spatial variables explained a significant amount of variation in community structure, a substantial amount of stochasticity also affects these communities, probably associated with unpredictable colonisation and persistence of the plants.     

The relationship between respiration and temperature in leaves of the arctic plant Saxifraga cernua

Abstract Saxifraga cernua, a perennial herb distributed throughout the arctic and subarctic regions, shows high levels of dark respiration. The amount of respiration exhibited by leaves and whole plants at any temperature is influenced by the pretreatment temperature. Plants grown at 10°C typically show higher dark respiration rates than plants grown at 20°C. The levels of alternative-pathway respiration (or cyanide-insensitive respiration) in leaves of S. cernua grown at high and low temperatures were assessed by treating leaf discs with 0.25 mol m^?3 salicylhydroxamic acid during measurements of oxygen consumption. Alternative pathway respiration accounted for up to 75% of the total respiration. Tissues from 20°C-grown plants yielded a Q₁₀ of 3.37 for normal respiration, and of 0.97 for alternative-pathway respiration. Tissues from 10°C-grown plants yielded a Q₁₀ of 2.55 for normal respiration, and of 0.79 for alternative-pathway respiration. The alternative pathway does not appear to be as temperature sensitive as the normal cytochrome pathway. A simple energy model was used to predict the temperature gain expected from these high rates of alternative-pathway respiration. The model shows that less than 0.02°C can be gained by leaves experiencing these high respiration rates.     

Timing of sexual reproduction and reproductive success in the high-mountain plant Saxifraga bryoides L

SAXIFRAGA BRYOIDES L. is one of the plant species reaching the upper limits of distribution for flowering plants in the European Alps. Because of its abundance in the subnival and nival zones, we expected S. BRYOIDES to reproduce efficiently in the highly stochastic climate at higher altitudes. Investigations were carried out at two subnival sites (2650 m and 2880 m a.s.l.) in the Austrian Alps. We studied flowering phenology, dynamics of seed development, and reproductive success in the climatically different years from 2001 - 2004. For a nival plant species, S. BRYOIDES showed a particularly long prefloration period (6 - 9 weeks). From onset of anthesis until seed maturity took an individual flower another 6 - 7 weeks and all individuals at a site 9 - 10 weeks. The length of the prefloration period and seed histogenesis was temperature-dependent, whereas seed maturation seemed to be endogenously controlled. Only in the exceptionally long and warm growing season of 2003 did all fruits mature at a site. In the other years, the onset of winter conditions halted development in many fruits before maturity. The seed/ovule ratio of mature fruits was around 0.7 in all years. The relative reproductive success (RRS) ranged from zero to 0.7, depending on the site and year. In conclusion, S. BRYOIDES needs an unexpectedly long time to undergo reproductive development. Though fruit maturation is uncertain, the high S/O ratio of single intact fruits results in at least a small seed crop in most years. This seems to be sufficient to assure the spread and maintenance of S. BRYOIDES at higher altitudes. As a seed-risk strategist (Molau,1993), S. BRYOIDES would clearly benefit from a prolonged growing season, which might occur more often if climate warming continues.     

Comparative germination ecology of two altitudinal vicariant Saxifraga species endemic to the north of Spain

Seeds of high‐mountain species are thought to germinate rapidly, synchronously and at high percentages after a cold period, with limited dependence on the external environment; yet, empirical evidence only partially supports this behaviour. We performed a comparative study of the germination response of two closely related taxa along an altitude gradient in northern Spain. Seeds from several maternal families of six populations of Saxifraga trifurcata (lowland species) and S. canaliculata (highland species) were subjected to temperature and stratification treatments. Germination percentages and germination rates were analysed using generalised linear mixed modelling and accelerated failure‐time modelling. We found that germination percentages and germination rates were high and dependent on incubation temperature in both species. Within species, seeds from higher altitudes had higher germination percentages under all conditions. Cold–wet stratification negatively affected germination success, particularly in the lowland species. Overall, the highland species was less responsive to the experimental treatments and showed more synchronous germination patterns. We conclude that seeds from these two Saxifraga species germinate as efficiently as species from other habitats, but have a narrower germination response, probably due to the stronger selective pressures in their harsh environments. Finally, a cold, wet stratification period is not a prerequisite for the germination of high‐mountain S. canaliculata, and its strong negative effect on the germination of its lowland relative S. trifurcata may contribute to the altitudinal segregation of these two species.     

Seed set and germination in crosses within and between two geographically isolated small populations of Saxifraga hirculus in Denmark

Dahlgaard, J. & Warncke, E. 1995. Seed set and germination in crosses within and between two geographically isolated small populations of Saxifraga hirculus in Denmark. - Nord. J. Bot. 15: 337–341. Copenhagen. ISSN 0107–055X.
Seed set and germination were studied in two geographically isolated populations of Saxifraga hirculus in Jutland, Denmark. The size of the populations differed, but both were assumed to exhibit significant genetic homogeneity and inbreeding due to (1) small population size (2) limited pollen flow and seed dispersal and (3) absence of gene flow between populations. Crosses within populations, as well as reciprocal crosses between populations, were carried out in the field. Seed number, proportion of developing seeds, germination percent, and germination rate were found to be greater for crosses between populations than for crosses within populations. Variation in seed size was found to be larger for maternal plants from the smaller population.     

Subtle differences in environmental stress along a flooding gradient affect the importance of inter-specific competition in an annual plant community

Empirical evidence suggests that the direction and intensity of plant–plant interactions may depend on the favourability of the environment. Previous studies have mainly focused on steep gradients of environmental stress or disturbance, while the interplay of competition and environment has not been tested for subtle environmental differences. Here, we present results from a study on plant communities of temporary wetlands in East-German farmland. Due to yearly ploughing in autumn, the vegetation is composed of annual species. Flooding does not affect adult plants and the elevation on the gradient expresses differences in the length of the growing season rather than in disturbance intensity or severe environmental stress. We tested whether such subtle differences in environmental stress may affect the importance of interspecific competition by the dominant species. Two treatments were applied at two elevations: removal of the dominant species (Matricaria maritima ssp. inodora) and reciprocal transplants of the seed-bank of the two elevations. At both elevations, removal of Matricaria inodora led to an increase in total species richness and number of wetland species, but the effects were substantially stronger at high elevations. Removal and the elevation on the flooding gradient significantly influenced the plant community composition. In particular, the weed communities became more similar to the wetland communities after the removal. Transplanted weed species did not emerge at low elevations. While two of four target species had significantly higher densities after the removal at high elevations, none of them was influenced by removal at low elevations. This indicates that, consistent with previous studies from other habitat types, competition by the dominant species was more intense under conditions of low environmental stress. The overall results suggest that both flooding as well as interspecific competition are important in structuring the plant communities along the freshwater gradient studied.     

Role of temperature in the germination ecology of three summer annual weeds

Summary Ambrosia artemisiifolia L., Chenopodium album L., and Amaranthus retroflexus L. are three summer annual weeds that occur in disturbed habitats. In nature, the peak germination season for A. artemisiifolia and C. album is in early to mid-spring, while in A. retroflexus the peak germination season is late spring to early summer. Furthermore, seeds of A. artemisiifolia germinate only in spring, while seeds of C. album and A. retroflexus germinate throughout the summer. In an attempt to explain the differential germination behavior of these three species in nature, changes in their germination responses to temperature during burial in a non-heated greenhouse from October 1974 to October 1975 were monitored. A high percentage of the seeds of all three species after-ripened during winter. Seeds of A. artemisiifolia and C. album germinated at temperatures characteristic of those in the field in early and mid-spring, but seeds of A. retroflexus required the higher temperatures of late spring and early summer for germination. Seeds of all three species germinated to higher percentages in light than in darkness. Non-dormant seeds of A. artemisiifolia that did not germinate in spring entered secondary dormancy. On the other hand, seeds of C. album and A. retroflexus that did not germinate when temperatures first became favorable for germination, did not enter secondary dormancy and, thus, retained the ability to germinate at summer field temperatures during summer. Thus, temporal differences in the germination behavior of these three species are caused by the differential reaction of the seeds to temperature during the annual temperature cycle.     

三峡库区9种植物种子萌发特性及其在植被恢复中的意义

三峡大坝蓄水后形成的库区消涨带面临植被消亡、生态退化等问题。为了筛选出适用于库区消涨带植被恢复的植物, 将9种1年生植物种子放置在库区消涨带不同海拔进行水淹(W 165-8 m, 121 d;W 155-18 m, 230 d;W 147-26 m, 271 d), 然后在实验室条件下进行萌发, 研究在消涨带淹水胁迫下这些种子的萌发特性。结果表明: (1) 除马唐(Digitaria sanguinalis)、小蓬草(Conyza canadensis)、金色狗尾草(Setaria glauca)种子在各条件下萌发率都较低外, 不同水淹条件对萌发率影响不同: 与新鲜种子相比, W 165水淹后, 旱稗(Echinochloa hispidula)和婆婆针(Bidens bipinnata)种子萌发率显著上升, 其余种子萌发率均显著下降; W 155水淹后, 所有种子的萌发率都显著下降且只有鱧肠(Eclipta prostrate)、黄花蒿(Artemisia annua)、合萌(Aeschynomene indica)3个物种有萌发, 萌发率分别为11.0%、7.3%和2.7%; W 147水淹后, 旱稗和婆婆针种子萌发率显著上升, 鱧肠种子无显著差异, 其余种子萌发率显著下降。(2) 鱧肠、黄花蒿、婆婆针和旱稗种子比其它物种更耐水淹。W 165水淹后, 鱧肠、黄花蒿、婆婆针、旱稗种子萌发率分别为44.7%、42%、20.7%和4.3%, W 147水淹后分别为76.3%、23%、15%和26.3%, 高于其他物种。(3) 水淹后种子萌发时间格局也受到影响, 大部分种子起始萌发时间推迟、萌发速度变慢。鱧肠、黄花蒿、婆婆针和旱稗的种子对三峡库区消涨带的水淹胁迫具有一定的适应能力, 可根据它们对水淹条件适应能力的差异在消涨带不同海拔高度进行植被恢复。     

温带半干旱地区一年生植物种子的萌发特性

在实验室条件下研究了中国温带半干旱地区科尔沁沙地的23种1年生植物的种子萌发特性（新采集种子、冷藏和干藏种子）.大籽蒿、虎尾草、冠芒草、沙蓬和地锦的新种子萌发率达90%左右,11种植物新种子萌发率均低于70%,说明这些植物的新种子具有或多或少的休眠属性.经过150d的冷干藏后,大籽蒿、虎尾草萌发率保持在90%以上,说明这两种植物完全没有休眠机制;冠芒草、沙蓬和地锦的种子萌发率下降较多,可能是储藏的环境条件导致的2次休眠现象;冷藏和干藏处理均能使绿珠藜、毛马唐、细叶益母草、雾冰藜、金狗尾草、苋菜、马齿苋、碱地肤和水稗草的种子在生长季开始时完成生理后熟,萌发率达到80%以上;干藏有利于促进毛马唐、细叶益母草、马齿苋和鹤虱的种子成熟,冷藏有利于促进绿珠藜和金狗尾草的种子成熟;黄蒿、灰绿藜、画眉草和烛台虫实在不同处理下的萌发率都比较低,说明种子内在生理休眠作用较强,具有减少种子一次性萌发数量的风险分摊策略.大多数1年生植物均能在较短时间内达到最终萌发率的90%,表现出迅速萌发的特性;黄蒿、灰绿藜、碱地肤和沙蓬种子则在不同处理中表现出延长萌发时间的策略来适应半干旱地区不确定的环境条件.最后,探讨了几种主要1年生植物的种子萌发对策与其对环境适应机制之间的关系.