Flowering Phenology and Gender Variation in Pennisetum typhoides

Because of the modular structure of pearl millet (an annual grass crop, Poaceae), different tillers of a plant share the same genotype but are subjected to different environmental conditions during their maturation. This allows investigation of the effects of tiller flowering phenology on allocation to resource-producing photosynthetic biomass, sexual functions, and thus tiller gender. All tillers of plants of two families collected from individual maternal plants (represented by 33 and 31 plants each) were analyzed. In both families, allocation to aboveground vegetative biomass decreased as flowering was delayed. On average, late-flowering tillers were 65% smaller than the first ones to flower. The proportion of biomass allocated to reproduction significantly increased with the flowering rank of the tillers, suggesting that translocations of assimilates occurred between early- and late-flowering tillers. In both families, late-flowering tillers produced significantly fewer pollen grains per stamen than early-flowering ones, and female reproductive allocation (expressed as seed mass per tiller) was also affected by flowering phenology. Tillers became increasingly female as flowering phenology progressed. This gender variation is possibly adaptive because pollination efficiency is maximized by plant height. Natural selection may favor a shift toward femaleness to maximize reproductive fitness in small, late-developing tillers.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Changes in reproductive investment with altitude in an alpine plant

Aims In perennial species, the allocation of resources to reproduction results in a reduction of allocation to vegetative growth and, therefore, impacts future reproductive success. As a consequence, variation in this trade-off is among the most important driving forces in the life-history evolution of perennial plants and can lead to locally adapted genotypes. In addition to genetic variation, phenotypic plasticity might also contribute to local adaptation of plants to local conditions by mediating changes in reproductive allocation. Knowledge on the importance of genetic and environmental effects on the trade-off between reproduction and vegetative growth is therefore essential to understand how plants may respond to environmental changes.Methods We conducted a transplant experiment along an altitudinal gradient from 425 to 1?921 m in the front range of the Western Alps of Switzerland to assess the influence of both altitudinal origin of populations and altitude of growing site on growth, reproductive investment and local adaptation in Poa alpina .Important findings In our study, the investment in reproduction increased with plant size. Plant growth and the relative importance of reproductive investment decreased in populations originating from higher altitudes compared to populations originating from lower altitudes. The changes in reproductive investment were mainly explained by differences in plant size. In contrast to genetic effects, phenotypic plasticity of all traits measured was low and not related to altitude. As a result, the population from the lowest altitude of origin performed best at all sites. Our results indicate that in P. alpina genetic differences in growth and reproductive investment are related to local conditions affecting growth, i.e. interspecific competition and soil moisture content.     

<Emphasis Type="BoldItalic">r</Emphasis>-Selected Traits in an Invasive Population

I compared life-history traits and self-fertilization rates in greenhouse culture of native Spartina alterniflora with an invasive population from a low-density Pacific estuary to see whether plants in the novel r-selective regime exhibit early reproduction, greater self-compatibility and high reproductive effort putting individuals at greater risk of death. Plants were grown from seed collected from the Atlantic and Gulf coasts of North America and a population introduced approximately 100 years ago to Willapa Bay, Washington, USA that has expanded to cover over ca. 6000 hectares. Pacific marshes have almost no native emergent vascular plants in the intertidal habitat, offering a virtually empty niche for invasive S. alterniflora. The low-density plants at the leading edge of this rapidly expanding population suffer a severe Allee effect of greatly reduced fecundity, caused by pollen limitation, compared to high-density areas. I found 99% of the invasive plants initiated reproduction in the first year of this study, while only 52% of the native range plants did so in the first year, followed by 34% in the second and 14% of native plants had not initiated reproduction after three years. The invasive plants had more than twice the reproductive effort of the native plants. Plants with the highest reproductive effort in the first year of growth died regardless of coastal site of origin, although nearly all of those that died over two years were invasive Pacific plants (27%). The invaders set two times the seed as the natives under forced selfing, suggesting greater self-compatibility or reproductive effort. These results suggest either a founding and bottleneck event and/or the invasive population has evolved from the long-lived, predominantly self-incompatible, K-selected state of the probable invasive propagules originating in dense, competitive native marshes.Co-ordinating editor: I. Olivieri     

Root proliferation characteristics of seven perennial arid-land grasses in nutrient-enriched microsites

We compared root proliferation in fertilized microsites among seven cultivars of five commonly planted cool-desert perennial grass species that differ in productivity and competitive ability. In a greenhouse experiment on nutrient-limited plants, one soil microsite in each pot received distilled water (control) and a second microsite received a rich, complete nutrient solution (fertilized). Roots in and adjacent to the microsites were mapped on Mylar windows for 22 days after the injections to determine the magnitude and timing of response in root length relative growth rates (RGRs). Because we provided adequate water, used a high level of fertilization in the treatment microsites, and conducted the experiments during rapid vegetative growth, the results provide a measure of the relative capacities and maximal rates of the grasses responses to enriched microsites. Root samples were harvested from control and fertilized microsites at the end of the experiment to determine the morphological basis of the proliferation response. In all seven grasses fine roots proliferated in the fertilized microsites faster than in the control microsites. The grasses did not differ in the timing of their response which showed a peak 7–8 days after injection. Although one species, Pseudoroegneria spicata cv. Goldar, had higher maximum root length RGR and higher RGR ratio (RGR in fertilized to RGR in control microsites) 7–8 days after injection, the seven grasses did not differ significantly in the magnitude of root length RGR response to fertilizer integrated over the 22 day experiment. The grasses also did not differ significantly in root morphological changes in fertilized mocrosites. Compared to roots in control microsites, roots in fertilized microsites had greater specific root length, length of secondary roots per length of main axis, number of lateral and sublateral roots per length of main axis, and mean lateral root length. Root proliferation was mainly the result of increased lateral branching and lateral root growth in all seven grasses. The consistency of root proliferation responses among these seven cultivars suggests that differences in the capacity for, maximum rate, or morphological basis of root proliferation are not directly related to ecological characteristics such as productivity and competitive ability. Other aspects of root response to nutrient enrichment, such as differential responses as a function of microsite nutrient concentration, plant phenology, plant nutrient status, or specific nutrient element(s), may still be important, but further experiments are required to determine whether different responses to enriched soil microsites among species correspond with know species differences in ecological characteristics.     

Water availability and population origin affect the expression of the tradeoff between reproduction and growth in Plantago coronopus

Investment in reproduction and growth represent a classic tradeoff with implication for life history evolution. The local environment can play a major role in the magnitude and evolutionary consequences of such a tradeoff. Here, we examined the investment in reproductive and vegetative tissue in 40 maternal half‐sib families from four different populations of the herb Plantago coronopus growing in either a dry or wet greenhouse environment. Plants originated from populations with an annual or a perennial life form, with annuals prevailing in drier habitats with greater seasonal variation in both temperature and precipitation. We found that water availability affected the expression of the tradeoff (both phenotypic and genetic) between reproduction and growth, being most accentuated under dry condition. However, populations responded very differently to water treatments. Plants from annual populations showed a similar response to drought condition with little variation among maternal families, suggesting a history of selection favouring genotypes with high allocation to reproduction when water availability is low. Plants from annual populations also expressed the highest level of plasticity. For the perennial populations, one showed a large variation among maternal families in resource allocation and expressed significant negative genetic correlations between reproductive and vegetative biomass under drought. The other perennial population showed less variation in response to treatment and had trait values similar to those of the annuals, although it was significantly less plastic. We stress the importance of considering intraspecific variation in response to environmental change such as drought, as conspecific plants exhibited very different abilities and strategies to respond to high versus low water availability even among geographically close populations.     

Population differentiation in the tiller architecture of Microstegium vimineum (Poaceae) in relation to habitat

Grass tillers grow by addition of modular units known as phytomers. Differences in phytomer organ size produce subindividual variation with potential adaptive value. Here, patterns in organ mass along tillers in the invasive annual Microstegium vimineum are related to habitat and tiller architecture. In an earlier study, seed families were collected from two populations: one from a sunny, woodland edge and the other from a shady understory in New Jersey, USA. Plants from these seeds were grown in a greenhouse. Phytomers along primary tillers were divided into culms, leaves, and cleistogamous or chasmogamous spikelets and seeds, dried and weighed. These data were used to examine the quantitative genetics of subindividual variation among families and populations, and the relationship of tiller fitness (based on total seed mass) to the mean or subindividual variance of phytomer traits. Phytomer position along a tiller was the major determinant of organ mass. Leaf mass increased from basal to upper nodes; cleistogamous reproductive mass decreased from upper to lower nodes. Phytomer organs were heaviest in the population from the sunny habitat. Family explained < 18% of variation in organ mass. Tiller fitness was positively correlated with mean culm, and leaf mass, but negatively correlated with coefficients of variation. Field‐collected tillers showed evidence of selection for increased leaf mass. Subindividual variation in M. vimineum is mostly due to phytomer position along a tiller and the prevailing light environment. Differentiation between sunny and shady populations suggests selection favors heavier phytomer leaves and culms, especially in the shady understory where this species is most invasive.     

A modular approach to biomass allocation in an invasive annual (Microstegium vimineum; Poaceae)

The invasive grass Microstegium vimineum grows in low light beneath the canopy of eastern forests in North America by reiteration of modules (phytomers) along a tiller. Basal phytomers are vegetative; terminal phytomers produce a raceme of chasmogamous (CH) spikelets plus an axillary raceme of cleistogamous (CL) spikelets. Additional subterminal phytomers with CL racemes mature basipetally. Allocation to culms, leaves, and CH and CL within phytomers was examined in relation to light conditions for a population in New Jersey, USA. Plants were reared in a greenhouse from seed families of parents in deep shade (2-8% full sun) or sunny, edge habitats. Primary tillers were subdivided into phytomers, dried, and weighed. Tillers from field habitats were similarly treated. For vegetative and subterminal phytomers, allocation to leaves and CH was greatest for the shady habitat. CL allocation decreased from terminal to reproductively immature subterminal phytomers. CH and CL mass was positively correlated with leaf mass, suggesting that reproduction is determined by available photosynthate. CH mass showed a genetic correlation with leaf mass. Developmental plasticity in modular allocation allows Microstegium to maximize fitness when conditions are favorable (e.g., high light along forest edges) by continual maturation of CL caryopses on axillary racemes.     

THE COST OF MERISTEM LIMITATION IN POLYGONUM ARENASTRUM: NEGATIVE GENETIC CORRELATIONS BETWEEN FECUNDITY AND GROWTH

Growth and reproduction in higher plants depend on meristems, which have three developmental fates. A meristem can become reproductive, but doing so terminates its activity, it can differentiate vegetatively, or it can remain quiescent for extended periods. The first two fates are mutually exclusive, and only the second leads to the production of additional meristems for subsequent growth and reproduction. In Polygonum arenastrum (frequently referred to as P. aviculare in North American Floras), an annual species lacking quiescent meristems, a quantitative genetic analysis of inbred full-sibling families revealed genetic variation in the developmental pattern of axillary meristem commitment to vegetative growth versus reproduction. Developmental variation resulted in family differences in the age of first reproduction, in age-specific fecundity and growth, and in final plant size and reproductive output. Furthermore, there were strong negative genetic correlations between age-specific growth and fecundity. Early commitment of meristems to reproduction favors high early fecundity, but reduces the number of meristems available for vegetative differentiation, and leads to lowered growth rates and fecundity later in life, when meristems are limiting. Conversely, meristem commitment to vegetative growth early in life results in low early fecundity but high late fecundity and growth. Meristem limitation, like resource limitation, is a proximate mechanism that generates trade-offs between life history traits. Differences between meristem limitation and resource limitation are discussed. Meristem limitation leads automatically to a senescent life history because of the determinate fate of reproductive meristems. Developmental characters were also found to be genetically correlated with metamer characters (leaf size, internode length) and seed size in this selfing species. The pattern of correlation is suggestive of selection for particular suites of life history and morphological characters.     

Spatial and seasonal patterns of microsite light availability in a remnant fragment of deciduous riparian forest and their implication in the conservation ofArisaema heterophyllum, a threatened plant species

Seasonal and spatial patterns of light availability were investigated in the understory of a small fragment (approximately 1300 m²) of a riparian deciduous forest of the Kokai River in central Japan dominated byQuercus acutissima Carruth., with the aim to understand the characteristics of microsite light availability forArisaema heterophyllum Blume, a threatened plant species uniquely associated with the riparian habitat. Diffuse site factor, which is the ratio of PFD (photon flux density, 400–700 nm) at a microsite to the open sky reference under diffuse light condition, was shown to be a satisfactory index for the evaluation of light availability for the understory plants in the habitat. Diffuse site factor 1,0 1,000 understory microsites along a 20 m transect from the edge to the interior of the forest, showed conspicuous seasonal changes in both mean and variation. Light availability decreased with seasonal tree canopy regeneration, with the highest spatial heterogeneity being recorded during the time of canopy closure. Auto-correlations of microsite light availability between different seasons were considerably high, suggesting the stability of relative light availability for individual microsites throughout the growing season ofA. heterophyllum. Fairly high light availability during summer season, which surpassed 20% of the open sky reference in most microsites, would be important for the growth and persistence of summer herbaceous plants likeA. heterophyllum.     

COST OF REPRODUCTION IN POLEMONIUM VISCOSUM: PHENOTYPIC AND GENETIC APPROACHES

I measured the effect of early reproduction on subsequent growth and survival in the alpine perennial wildflower, Polemonium viscosum. Measurements were made over 4 yr on 34 maternal sibships under natural conditions. A significant phenotypic cost of early reproduction characterized the study population. Plants that flowered after only one year's growth had twice as many leaves and 25% more shoots than nonflowering individuals of equal age. However, early flowering decreased leaf number by 18% in the subsequent year and survivorship by 20% after two years relative to changes in leaf number and survival of nonflowering plants. For such trade-offs to shape the further evolution of reproductive schedules, flowering probability and those age-specific components of plant size that represent the energetic currency for reproductive costs must be heritable. Although families showed significant heterogeneity in the probability of early flowering, most (62%) entirely failed to flower. Moreover, phenotypic variation in vegetative size components at ages 1 and 2 had little genetic basis. Only at ages 3 and 4, after vegetative and demographic costs of early reproduction had been incurred, did vegetative size components (leaf length and number, and shoot number) vary significantly among families. Results of this study provide little evidence of a genetically based trade-off between early reproduction and subsequent survival in P. viscosum.     

Habitat and microsite influence demography of two herbs in intact and degraded scrub

Identifying environmental factors associated with vital rate variation is critical to predict population consequences of environmental perturbation. We used matrix models to explore effects of habitat and microsite on demography of two widespread herbs, Chamaecrista fasciculata (partridge pea) and Balduina angustifolia (yellow buttons). We evaluated models simulating population dynamics in common microsites (shrub, litter, bare sand) within two habitats (intact, degraded Florida scrub) using data on experimental populations initiated by sowing seeds, and natural seed production. Models included four stages (seed bank, small vegetative, large vegetative, reproductive) and three vital rates (survival, growth, fecundity), summarized in sixteen transitions. We conducted life table response experiments to assess contributions of each habitat and microsite to population growth rates. We found that (1) C. fasciculata had greatest population growth in degraded habitat and litter microsites, (2) B. angustifolia had similar population growth between habitats and greatest in bare sand microsites, (3) advancing growth transitions of C. fasciculata had greatest elasticity on population growth in degraded habitat, shrub, and litter, as did seed survival in intact habitat and bare sand, (4) seed survival and advancing growth transitions of B. angustifolia had greatest elasticity on population growth in both habitats, as did seed survival in shrub and litter, and advancing growth in bare sand. Greater population growth of C. fasciculata in degraded scrub is probably explained by release from belowground competition; B. angustifolia may be most affected by competition with shrubs. Microsites in intact scrub were not ecologically equivalent to those in degraded scrub emphasizing that intact scrub is ecologically complex and critical to preserve.     

Resource allocation to sexual and vegetative reproduction in a forest herbSyneilesis palmata (Compositae)

The balance between sexual and vegetative reproduction inSyneilesis palmata was examined in relation to environmental conditions and the amount of reproductive resources, which is defined here as the total quantity of dry matter invested in both modes of reproduction. The allocation balance was measured for individual plants of two populations, with different densities in an open habitat, for 2 years, and those of two other populations, under different light intensities in a plantation forest (forest floor and edge), for 3 years. Relative allocation to sexual reproduction decreased with increasing reproductive resources in all populations except for the forest edge, which showed a constant allocation balance. The high density population showed lower relative allocation to sexual reproduction than the low density population, irrespective of the amount of reproductive resources. However the between-year comparison of the high density population suggested that under extremely high density, plants with a small amount of reproductive resources enhanced sexual reproduction, while plants with a large amount of reproductive resources reproduced vegetatively. On the forest floor, plants with small amounts of reproductive resources had higher relative allocation to sexual reproduction than the forest edge population, while plants with large amounts of reproductive resources had a somewhat lower one. The adaptive significance of such allocation patterns are discussed, based on qualitative data on the characteristics of both types of offspring.     

Effects of competitive stress on vegetative growth, storage, and regrowth after defoliation in Phleum pratense

The capacity for resource storage is important to population persistence in nonforaging plants with a consolidation strategy. In a competitive environment, selection may favor genetic variants with a well-developed ability to store resources for future vegetative growth or to regrow following herbivory. To determine the evolutionary potential for changes in vegetative growth, storage and regrowth after defoliation in response to competitive stress, half-sib families of the caespitose grass Phleum pratense were grown in pots in a glasshouse either alone or in competition with four individuals of Lolium perenne . They were defoliated after 16 wk and permitted to regrow for 8 wk. Production of new tillers, leaf area and leaf dry mass were recorded before and after defoliation; resource storage was estimated by the dry mass of stem bases. Tiller numbers, leaf area and mass, regrowth, and stem base mass were significantly reduced by competition; however, there was significant variation among families for many variables. A significant proportion of the variation in clipped and regrowth leaf mass, and in stem base mass, was due to a competition by family interaction. Storage in stem bases was correlated with total accumulated leaf mass in both control and competition groups. Genetic variation for competitive ability and the ability to regrow rapidly after defoliation exist in this P. pratense population. A well-developed ability to regrow following release from competitive or grazing pressures is likely to be a critical adaptation of caespitose grasses.     

Subdominant species distribution in microsites around two life forms at a desert grassland‐shrubland transition zone

Question: In the same landscape context — at a desert grassland‐shrubland transition zone, how does subdominant plant abundance vary in microsites around dominant grasses and shrubs? Location: Sevilleta LTER, New Mexico, USA (34°21’N; 106°53’W; 1650 m a.s.l.). Methods: We compared the distribution of subdominant plants in canopy, canopy edge and interspace microsites around individual shrubs (Larrea tridentata) and grasses (Bouteloua eriopoda) at a transition zone that has been encroached by shrubs within the past 50 ‐ 100 a. Plots of variable size according to microsite type and dominant plant size were sampled. Results: Subdominant abundance was higher in microsites around L. tridentata shrubs than in microsites around B. eriopoda. Furthermore, differences in species abundance and composition were higher among microsites around grasses than among microsites around shrubs. The distribution of subdominants was mostly explained by their phenological characteristics, which indicates the importance of temporal variation in resources to their persistence. Conclusions: This study of coexistence patterns around dominants revealed ecological contrasts between two dominant life forms, but other factors (such as disturbances) have to be taken into consideration to evaluate landscape‐scale diversity.     

The enemy release and EICA hypothesis revisited: incorporating the fundamental difference between specialist and generalist herbivores

The success of invasive plants has been attributed to their escape from natural enemies and subsequent evolutionary change in allocation from defence to growth and reproduction. In common garden experiments with Senecio jacobaea, a noxious invasive weed almost worldwide, the invasive populations from North America, Australia, and New Zealand did indeed allocate more resources to vegetative and reproductive biomass. However, invasive plants did not show a complete change in allocation from defence to growth and reproduction. Protection against generalist herbivores increased in invasive populations and pyrrolizidine alkaloids, their main anti‐herbivore compounds, did not decline in invasive populations but were higher overall compared with native populations. In contrast, invasive plants lost additional protection against specialist herbivores adapted to pyrrolizidine alkaloids. Hence, the absence of specialist herbivores in invasive populations resulted in the evolution of lower protection against specialists and increased growth and reproduction, but also allowed a shift towards higher protection against generalist herbivores.     

横断山区干旱河谷川滇蔷薇生长与繁殖特征的地理空间差异

通过大面积的野外踏查、种群调查和室内实验,分析了横断山区干旱河谷乡土灌木川滇蔷薇在岷江流域、大渡河流域、雅砻江流域和金沙江流域的分布、生长繁殖特征及其在流域间的差异。结果表明,川滇蔷薇广泛分布于四大流域,海拔范围1700m至3700 m,所调查的18个种群生长状况良好,株高2 m以上,明显高于群落中的其他灌木。主成分分析表明,地理位置(流域、纬度、经度、海拔)和土壤(全碳、总有机碳、全氮、pH值)是影响川滇蔷薇生长繁殖的主要因子,其中地理位置的影响更大。生长特征包括株高、地径和冠幅在18个种群间差异显著,但在不同的流域间差异不显著。丛萌生数在种群间差异显著,其差异性较其他生长特征弱,但丛萌生数在不同流域间差异显著:岷江流域显著大于雅砻江流域,金沙江流域显著大于大渡河流域。与生长和无性繁殖相比,有性繁殖特征表现出更强的空间差异性,果实重量、瘦果重量以及种子活力在居群间差异极显著,果实重量和种子活力在不同流域间差异显著,但瘦果重量在流域间的差异不显著。综合分析表明,川滇蔷薇生长和繁殖特征在横断山区干旱河谷存在较强空间差异性,较强无性繁殖是种群持续存活的保证,而有性繁殖及其强烈的空间差异为种群适应多变环境以及地理空间扩展提供了遗传基础。     

Resistance and tolerance in a host plant-holoparasitic plant interaction: genetic variation and costs

Host organisms are believed to evolve defense mechanisms (i.e., resistance and/or tolerance) under selective pressures exerted by natural enemies. A prerequisite for the evolution of resistance and tolerance is the existence of genetic variation in these traits for natural selection to act. However, selection for resistance and/or tolerance may be constrained by negative genetic correlations with other traits that affect host fitness. We studied genetic variation in resistance and tolerance against parasitic infection and the potential fitness costs associated with these traits using a novel study system, namely the interaction between a flowering plant and a parasitic plant. In this system, parasitic infection has significant negative effects on host growth and reproduction and may thus act as a selective agent. We conducted a greenhouse experiment in which we grew host plants, Urtica dioica, that originated from a single natural population and represented 20 maternal families either uninfected or infected with the holoparasitic dodder, Cuscuta europaea. that originated from the same site. We calculated correlations among resistance, tolerance, and host performance to test for costs of resistance and tolerance. We measured resistance as parasite performance (quantitative resistance) and tolerance as the slopes of regressions relating the vegetative and reproductive biomass of host plants to damage level (measured as parasite biomass). We observed significant differences among host families in parasite resistance and in parasite tolerance in terms of reproductive biomass, a result that suggests genetic variation in these traits. Furthermore, we found differences in resistance and tolerance between female and male host plants. In addition, the correlations indicate costs of resistance in terms of host growth and reproduction and costs of tolerance in terms of host reproduction. Our results thus indicate that host tolerance and resistance can evolve as a response to infection by a parasitic plant and that costs of resistance and tolerance may be one factor maintaining genetic variation in these traits.     

Seasonal control over allocation to reproduction in a tussock-forming and a rhizomatous species of Eriophorum in central Alaska

Summary The evergreen tussock-forming Eriophorum vaginatum revealed consistently earlier (c. 1 moth) phenology and greater biomass per tiller than the summergreen rhizomatous E. scheuchzeri in all four components measured (vegetative and reproductive shoots and stems) under the same climatic regime in central Alaska over one growing season. Greatest allocation to vegetative shoot growth occurred in mid-summer in both species. The tussock growth form of E. vaginatum raised shoot meristems 25–30 cm above the soil surface, where temperatures were warmer, permitting shoot growth to begin earlier in spring and continue longer in autumn than in E. scheuchzeri. Consequently, E. vaginatum was able to allocate reserves to reproductive tillers primarily in autumn and early spring, times when minimal reserves were required for vegetative growth. By contrast, the rhizomatous E. scheuchzeri had a more constrained growing season, and allocation to reproduction coincided with allocation to vegetative growth. For this reason, reserves were drawn down more fully in mid-summer in E. scheuchzeri than in E. vaginatum. The more conservative use of nutrient stores in E. vaginatum may relate to its great longevity, reduced allocation to reproduction (including low seedling recruitment), and relatively stable habitats. The mid-seasonal pulse of allocation to reproduction in E. scheuchzeri appears viable only in relatively fertile disturbed sites, where the soil nutrient supply is sufficient to support simultaneous allocation to vegetative growth and reproduction.     

Phenology and life cycle of the annual, <Emphasis Type="Italic">Chamaesyce maculata</Emphasis> (L.) Small (Euphorbiaceae), with multiple overlapping generations in Japan

The phenology of germination, vegetative growth and sexual reproduction in the annual Chamaesyce maculata (L.) Small (Euphorbiaceae) were investigated in a natural population in western Japan. Seedlings emerged from mid-June to early October, with three peaks: mid-June, late July and late August. Plants that emerged in June commenced sexual reproduction from late July, and thereafter both vegetative growth and sexual reproduction occurred together until early November, the plants showing no switching from vegetative growth to sexual reproduction. Seedlings that emerged in June and July suffered high mortality, but most seedlings that emerged from August onward survived until the reproductive stage. The minimum size for reproduction was largest for plants that emerged early in the season, and it decreased with a delay in seedling emergence. The late emergence of seedlings that resulted in low reproductive output may be to some extent compensated for by the increased probability of survival in the seedling stage. A transplant experiment clarified that C. maculata can repeat a maximum of three overlapping generations within a year. Multiple generations per year were attained by non-dormant seeds produced in the first and second generations and clearly resulted in an increased reproductive output per year. The life cycle with multiple overlapping generations may have been acquired in habitats where unpredictable disturbance results in temporally unsuitable conditions for germination, vegetative growth and sexual reproduction of annual plants, but where suitable conditions frequently continue over a period longer than the single generation time of annual plants.