Reproductive biology of a primitive angiosperm,Pseudowintera colorata (Winteraceae), and the evolution of pollination systems in theAnthophyta

Stigmatic exudate-eating chironomid flies (Smittia) and pollen-eating halodid beetles are the most common visitors to flowers ofPseudowintera colorata. Plants are self-incompatible. After selfing pollen tubes penetrate the nucellus; chase pollination experiments show that the incompatibility reaction occurs before fertilization. When pollen is applied to stigmas, the exudate dries rapidly and does not usually reappear. A review of anthophytes suggests that their common ancestor around 200 mya was bisexual and pollinated at least in part by nectar-seeking insects, possibly including flies. The angiosperm stigma appears to have evolved from a pollination drop mechanism via an adaxial stigmatic surface.Dedicated with pleasure to ProfessorW. R. Philipson on the occasion of his 80th birthday.     

深山含笑二次开合开花动态与雌雄异熟特征

对深山含笑（Michelia maudiae）的花部特征、开花动态、开花物候、花粉活力和柱头可授性进行研究。结果表明：①深山含笑花期为2月下旬至4月中上旬,表现为集中开花模式;②单花开放时间固定,且有二次开合现象,开放过程可分为露白、首次开花期、闭合期和二次展开期共4个时期;③深山含笑在单株水平上为同步集中式开花,单花花期约为7~12 d,单株花期约为20~38 d;④深山含笑柱头在开花前1 d已具有可授性,开放当天可授性最高,随后可授性迅速下降,而雄蕊在开花第2 d开始散粉,花粉活力最高时期为开花第3 d,为（64.6±6.1）%;⑤深山含笑为雌雄异熟且雌蕊先熟,并且雌雄特征有重叠表达阶段,这样不仅可有效减少自交,又可支持自交,足以保证生殖成功。     

Plastic bet-hedging in an amphicarpic annual: an integrated strategy under variable conditions

Amphicarpy is a form of diversified bet-hedging expressed mostly in annual plants, where two types of offspring are produced with two distinct ecological roles: long-range aerial dispersers and highly competitive subterranean, sedentary fruit. Emex spinosa is a semi-arid, amphicarpic annual, inhabiting habitats with different levels of environmental variation. We tested the hypothesis that, in E. spinosa, bet-hedging may be “fine-tuned” by plasticity in the phenotype ratio (aerial/subterranean fruit mass) as a function of environmental conditions. We conducted a greenhouse experiment, manipulating nutrient availability and intraspecific density, to determine the pattern of ratio shifts. In order to determine whether the integrated strategy is an adaptation to variable habitats, a similar common garden experiment was conducted, comparing two natural populations differing in environmental variability. The offspring ratio shifted in response to both nutrient availability and plant density. In pots containing single plants the ratio increased steeply with nutrient availability, while in pots containing eight plants a more moderate increase occurred. These shifts were the result of plasticity in allocation to both achene types, as well as ontogenetic effects on aerial achene production. The degree of response increased with the heterogeneity of the habitat of origin. We found evidence for an adaptive integrated strategy, with bet-hedging “fine-tuned” by phenotypic plasticity. Strenuous conditions tended to shift the offspring ratio towards securing subterranean reproductive success, while favorable conditions resulted in a shift towards dispersible achenes. The authors Asaf Sadeh and Hagai Guterman contributed equally to this study.     

Phenotypic gender variation within inflorescences of the protogynous species Helleborus foetidus L. (Ranunculaceae)

In the protogynous species H. foetidus, I investigated if investment in sexual structures and seed set shows any trend with position in the flower-opening sequence. In four-flower inflorescences, stamen number decreased with flower position i.e. was highest for the earliest flower and lowest for the latest flower. Ovule number was significantly higher in the latest flower. Stamen and ovule number did not covary, indicating that there is no structural gender trade-off in this species. Seed set (i.e. percentage of ovules producing seed) did not differ between control plants and pollen-supplemented plants and the effect of pollen supplementation did not vary among positions. Seed set showed marginally significant differences between control plants and simulated-herbivory plants, but the effects of this treatment varied significantly among positions. Significant among-position variation in seed set was observed in the control plants but not the simulated-herbivory plants. Seed set of latest-opening flowers of simulated-herbivory plants was significantly higher than that of latest-opening flowers of control plants. H. foetidus support Brunet and Charlesworth's (1995) prediction that in plants with protogynous flowers, later-opening flowers should specialize as female flowers, at least under conditions of high resource availability.     

Generic relationships and dating of lineages in Winteraceae based on nuclear (ITS) and plastid (rpS16 and psbA-trnH) sequence data

Phylogenetic analyses of representative species from the five genera of Winteraceae (Drimys, Pseudowintera, Takhtajania, Tasmannia, and Zygogynum s.l.) were performed using ITS nuclear sequences and a combined data-set of ITS + psbA-trnH + rpS16 sequences (sampling of 30 and 15 species, respectively). Indel informativity using simple gap coding or gaps as a fifth character was examined in both data-sets. Parsimony and Bayesian analyses support the monophyly of Drimys, Tasmannia, and Zygogynum s.l., but do not support the monophyly of Belliolum, Zygogynum s.s., and Bubbia. Within Drimys, the combined data-set recovers two subclades. Divergence time estimates suggest that the splitting between Drimys and its sister clade (Pseudowintera + Zygogynum s.l.) occurred around the end of the Cretaceous; in contrast, the divergence between the two subclades within Drimys is more recent (15.5–18.5 MY) and coincides in time with the Andean uplift. Estimates suggest that the earliest divergences within Winteraceae could have predated the first events of Gondwana fragmentation.     

Temporal variation in phenotypic gender and expected functional gender within and among individuals in an annual plant

Background and Aims

Adaptive explanations for variation in sex allocation centre on variation in resource status and variation in the mating environment. The latter can occur when dichogamy causes siring opportunity to vary across the flowering season. In this study, it is hypothesized that the widespread tendency towards declining fruit-set from first to last flowers on plants can similarly lead to a varying mating environment by causing a temporal shift in the quality (not quantity) of siring opportunities.

Methods

A numerical model was developed to examine the effects of declining fruit-set on the expected male versus female reproductive success (functional gender) of first and last flowers on plants, and of early- and late-flowering plants. Within- and among-plant temporal variation in pollen production, ovule production and fruit-set in 70 Brassica rapa plants was then characterized to determine if trends in male and female investment mirror expected trends in functional gender.

Key Results

Under a wide range of model conditions, functional femaleness decreased sharply in the last flowers on plants, and increased from early- to late-flowering plants in the population. In B. rapa, pollen production decreased more rapidly than ovule production from first to last flowers, leading to a within-plant increase in phenotypic femaleness. Among plants, ovule production decreased from early- to late-flowering plants, causing a temporal decrease in phenotypic femaleness.

Conclusions

The numerical model confirmed that declining fruit-set can drive temporal variation in functional gender, especially among plants. The discrepancy between observed trends in phenotypic gender in B. rapa and expected functional gender predicted by the numerical model does not rule out the possibility that male reproductive success decreases with later flowering onset. If so, plants may experience selection for early flowering through male fitness.     

Studying genetics of adaptive variation in model organisms: flowering time variation in Arabidopsis lyrata

Arabidopsis thaliana has emerged as a model organism for plant developmental genetics, but it is also now being widely used for population genetic studies. Outcrossing relatives of A. thaliana are likely to provide suitable additional or alternative species for studies of evolutionary and population genetics. We have examined patterns of adaptive flowering time variation in the outcrossing, perennial A. lyrata. In addition, we examine the distribution of variation at marker genes in populations form North America and Europe. The probability of flowering in this species differs between southern and northern populations. Northern populations are much less likely to flower in short than in long days. A significant daylength by region interaction shows that the northern and southern populations respond differently to the daylength. The timing of flowering also differs between populations, and is made shorter by long days, and in some populations, by vernalization. North American and European populations show consistent genetic differentiation over microsatellite and isozyme loci and alcohol dehydrogenase sequences. Thus, the patterns of variation are quite different from those in A. thaliana, where flowering time differences show little relationship to latitude of origin and the genealogical trees of accessions vary depending on the genomic region studied. The genetic architecture of adaptation can be compared in these species with different life histories.     

Flowering phenology of <Emphasis Type="Italic">Ulex europaeus</Emphasis>: ecological consequences of variation within and among populations

Reproductive phenology of gorse (Ulex europaeus L., Genisteae, Fabaceae) is unusual in that the onset and duration of flowering vary greatly among individuals within populations: some plants initiate flowering in autumn or winter and continue flowering through spring, others initiate flowering in early spring. To understand the origin of this diversity and its ecological consequences, we investigated flowering phenology of randomly sampled individuals from five different natural populations in Brittany (France). Reproductive success was evaluated for individuals with contrasting flowering patterns, from 16 natural populations. Flower production, pod production, seed production and seed predation were estimated. Plants initiating flowering in spring produced larger numbers of flowers and pods over a shorter period than plants flowering from winter to spring, which produced few flowers and pods at a time but over a longer period. Pod production of long-flowering plants did not differ significantly between winter and spring, but their pods were more intensively attacked by seed predators in spring than in winter. We discuss our results in relation to biotic and abiotic parameters. We postulate that long-flowering can be interpreted as a bet-hedging strategy, spreading the risk of pod failure (rotting or freezing) in winter and of seed predation in spring.     

Pollen flow,self-pollination and gender specialization: Factors affecting seed-set in the tristylous speciesLythrum salicaria (Lythraceae)

Some of the factors governing seed-set in three populations ofLythrum salicaria are examined during two seasons. Plants spatially segregated from legitimate neighbours by distances exceeding 1m set few seeds. Plants with small inflorescences set fewer seeds than those with medium or large inflorescences. Flowers having self pollen applied to the stigma before legitimate pollination set fewer seed per flower than those pollinated with legitimate pollen only. Seeds of the mid-style morph germinated more effectively than those of the long style morph and seeds of the short-style morph showed very low levels of germination.     

Life history variation and infraspecific heterochrony inRhinanthus glacialis (Scrophulariaceae)

Comparative cultivation experiments were made with 14 populations of the annual, hemiparasitic speciesRhinanthus glacialis from seven different habitat types. Phenological data were recorded and plants from field samples and from corresponding experimental samples were morphologically analysed.There are up to ten weeks difference in the start of flowering between means of the extreme experimental populations. Flowering time variation correlates with the number of internodes and also with most of the other morphological characters. Flowering time also shows close relationships to different grassland types and the examined populations appear to be well-adapted to the environmental conditions in natural or man-influenced grassland types as far as flowering time is concerned. The number of internodes is mostly insensitive to the applied experimental influences. The rate of internode production is constant. Trade-offs are suggested for some morphological traits (i.e., few internodes with few branches and thus few flowers as trade-off for early flowering). — Previous studies of life history variation based on population data have not been incorporated into the theory of heterochrony. The pattern of life history variation of populations ofR. glacialis together with information about morphology, ecology, and phylogeny suggests that some variants have evolved from ancestors via heterochronic processes. Earlier flowering populations from montane hay meadows, litter meadows, and dry meadows belong to paedomorphic variants. These have fewer internodes and branches as well as reduced height in comparison with the suggested ancestor. Later flowering populations from subalpine limestone grassland belong to a peramorphic variant with more internodes, more branches, and taller stems in comparison with the suggested ancestor.     

Variation and similarities in pollen features in some basal angiosperms, with some taxonomic implications

Species within three families of basal angiosperms (Trimeniaceae, Winteraceae, Monimiaceae) illustrate differences and similarities in pollen within a species, between species and between genera. Trimenia papuana (Trimeniaceae) has dimorphic pollen (inaperturate, polyforate), each confined to different individual plants. Other species have either disulculate or polyforate pollen. Evolution seems to be from disulculate to inaperturate to polyforate. Present-day Winteraceae have pollen in permanent tetrads except four species of Zygogynum with monads. Why? Did such monads appear as fossils before tetrads in Winteraceae? Molecular studies of Takhtajania perrieri indicate it is basal but its unique bicarpellate unilocular gynoecium seems derived. Although Hedycarya arborea and Kibaropsis caledonica have near-identical permanent pollen tetrads, many other features are very different. Hedycarya species have permanent tetrads or inaperturate monads with spinulose, `starry' or other sculpturing, and it is suggested this and recent molecular data indicate further studies are needed to determine generic limits.     

Size-dependent variation of gender in high density stands of the monoecious annual,Ambrosia artemisiifolia (Asteraceae)

Summary We examined the relationship between size variability and the distribution of functional gender in stands of the monoecious, wind-pollinated annual Ambrosia artemisiifolia. Populations of 60 individuals were grown in the greenhouse at a density of 372 m^–2 and at two nutrient levels. Among the surviving plants, after self-thinning, variability in above-ground biomass and gender was higher in the high nutrient treatment. Among individuals there was a significant positive correlation between maleness and both height and biomass. Fecundity was also positively correlated with both measures of size. Based on the pattern of distribution of male and female flowers within the plant, it appears that the increase in maleness in larger plants is due to increased branching and axis elongation. These results demonstrate that competitive interactions, which lead to increased variability in biomass and fecundity, can also generate variability in gender within populations.     

Intrapopulation gender variation in common ragweed (Asteraceae: Ambrosia artemisiifolia L.), a monoecious,annual herb

Summary We measured variation in gender among individuals within populations of ragweed (Ambrosia artemisiifolia) in an abandoned old-field and in the greenhouse. There was great variability in sex expression, from all-female to approximately 78% male. Plants differed significantly in gender in different locations within the field. Plants in an area abandoned from agriculture one year previously were more male than plants in a nearby area abandoned four years previously. In the greenhouse, soil moisture treatments and levels of attack by spittlebugs (Philaenus spumarius) did not affect gender. Height was positively correlated with relative maleness in both populations. Plants with greater shoot weight were relatively more male in the greenhouse, but not in the field. The gender variation we observed either has a genetic basis or is controlled by environmental variables other than those we investigated.     

Environmental variation and large-scale Gracilaria production

Temporal and spatial abiotic variation in seaweed farms should be anticipated to maximize production through alternative exploitation strategies. This study describes the basic assumptions and the most relevant data used to empirically develop a production model aimed at improving prediction and increasing production of Gracilaria farms in northern Chile. Continuous light and temperature recordings since 1986 have allowed us to relate abiotic variations with high production seasons of Gracilaria or with the presence of pests and epiphytes. Much of the interannual variations in light and temperature appear as part of a predictable pattern of change between ENSO (El Niño/Southern Oscillation) and inter-ENSO years. Production has been found to be a function of stock density and harvesting frequency, two parameters that can be effectively manipulated in the field. Thus, the range of climatic change now can be anticipated to some extent which, in turn, suggests the best farming strategy. During seasons or growth periods anticipated to be highly productive, farming activities are oriented to maintain high percentage removal of the stock through frequent harvesting. During seasons anticipated to be low in production, activities are oriented to prevent biomass losses due to the blooms of epiphytes and pests and to secure stocks to renew through planting the damaged parts of the beds after the blooms.     

Isozyme variation inFaidherbia albida (Leguminosae, Mimosoideae)

Genetic variation has been assessed in 30 populations (931 families) ofFaidherbia albida (Leguminosae, Mimosoideae) from across its entire African range, using six isozyme loci identified by five enzyme systems. Among the populations studied a null allele was proposed to explain the absence ofLap-1 activity in populations from southern and eastern Africa. The mean percentage of polymorphic loci per population, the mean number of alleles per locus and the mean genetic diversity within populations were 31.7%, 1.6 and 0.127 respectively. Genetic diversity was greatest in populations from West Africa and lowest in populations from eastern/southern Africa, with Ethiopian/Sudanese populations intermediate. The overall degree of genetic differentiation between populations (G_ST) indicated that approximately 56% of the enzyme variation resided within populations. Clustering of Nei's unbaised genetic distances calculated between all populations produced a dendrogram that generally followed the geographic distribution of the populations. Two major groups were identified that may be considered the eastern/southern African and the Ethiopian/West African clusters. Within the Ethiopian/West African cluster two subclusters could be recognised, one broadly corresponding to those populations from Ethiopia/Sudan and the other to those populations from West Africa. The implications of these results for theories regarding the origin ofF. albida in Africa are discussed.     

日龄及性别对橘小实蝇实验种群飞行能力的影响

利用飞行磨系统对实验种群橘小实蝇的1、6、11、16、21、26、31日龄的7个雌雄个体分别进行22h连续吊飞试验,测定其累计飞行距离、累计飞行时间、平均飞行速度。橘小实蝇原始种群采自湖南长沙,经室内连续饲养约30代,成为试验种群。试验环境条件为连续光照条件,温度25℃,RH 60%—80%,光照强度为990 lx。整个吊飞过程不补充食物和水份。共得到179组吊飞数据,其中雌虫86组,雄虫93组,每个日龄处理不少于9头。结果显示随着日龄的增加,橘小实蝇雌蝇和雄蝇的飞行能力均逐渐增加,到达16日龄后雌蝇飞行能力迅速下降,表现为单峰型;雄蝇的飞行能力在16—31日龄一般维持较高水平,但在26日龄有一定波动。16日龄雌蝇和雄蝇的累计飞行距离分别是(2485.6±2287.2)m和(2152.3±1773.3)m。其中1头21日龄雄性个体的累计飞行距离最大,完成了8795.80 m的飞行。雌蝇累计飞行距离最远的发生在16日龄,为8116.6 m。日龄对橘小实蝇的累计飞行距离和平均飞行速度的影响都达到了显著水平(前者,F=2.88,P0.05;后者,F=4.98,P0.01)。雌蝇和雄蝇的平均飞行速度的差异达到了显著水平(F=4.10,P0.05)。日龄和性别的互作对平均飞行速度的影响也达到了显著水平(F=2.74,P0.05)。尽管实验种群不能完全代表野生种群,研究结果对研究橘小实蝇扩散规律及综合防治,特别是改进橘小实蝇雄性不育技术仍有重要的参考价值。     

Flowering response of day-neutral and short-day cultivars of Nicotiana tabacum L. interactions among roots,genotype, leaf ontogenic position and growth conditions

The interaction between roots and leaves as a function of the capacity of differently positioned leaves to induce flowering of four cultivars of Nicotiana tabacum L. was assessed under long-and short-day growth conditions with three types of manipulations: 1) repeated rooting of the shoot tip, 2) removal of apical leaves, and 3) removal of basal leaves. Repeated rooting of the shoot tip increased the number of nodes produced by all cultivars; however, a substantial extension of vegetative growth was only caused by rerooting in conditions where apical leaves exhibited little or no inductive capacity. The simplest and most consistent interpretation of these data is that floral initiation in tobacco results from an interaction of inputs from the leaves and the roots and that the root influence can be overridden by a strong leaf signal.     

Somatic embryogenesis,plant regeneration and somaclonal variation in barley

In vitro culture of immature embryo and young leaf tissues was carried out with five cultivars of barley, Hordeum vulgare. Two cultivars (Albacete and Porthos) responded poorly from both types of explants, while the three others (Dissa, Golden Promise and Ingrid) produced a high frequency of embryogenic callus from these explants (25–60%). For Dissa and Ingrid, young leaf explants were slightly better than immature embryo explants for embryogenic callus induction, while immature embryo cultures of Golden Promise responded better than young leaf explants. Thus, there appears to be a significant genotype × explant interaction in the initiation of embryogenic callus in barley.Some phenotypic variants were detected among the regenerated plants of Golden Promise and Ingrid, most originating by epigenetic changes. Only in one case was the variant phenotype heritable, probably due to a mutation in the chloroplast DNA. Mitotic alteractions were not detected. Consequently, somaclonal variation did not appear to be a very frequent event in plants regenerated from 1- to 6- month-old cultures of barley.