POPULATION DYNAMIC CONSEQUENCES AND EVOLUTION OF SEED TRAITS OF HETEROSPERMA PINNATUM (ASTERACEAE)

We summarize research on variation in achene morphology of Heterosperma pinnatum. Each flowering head has a broad range of achene morphologies. There is no between-year seed bank. There are no ecologically relevant differences in achene size, seedling size, or seedling growth rates of the different morphs. Achenes located centrally in the heads lose dormancy earlier during the period between autumn achene production and the onset (May-July) of the rainy season. This results in relatively more early emergence of central achenes. Awned central achenes also have greater adhesive dispersibility. Emergence, survival, and fecundity were measured for achenes sown into natural populations in central Mexico. When harsh conditions result in few seeds surviving to reproduce, the advantage tends to go to the more “conservative” peripheral achenes. Greenhouse experiments show a strong genetic component to the determination of achene proportions within and among populations and a variety of genetic correlations. Populations with a high proportion of awned achenes tend to be found in vegetation types where they are likely to be ephemeral. A high proportion of central achenes tend to be found in sites with little pre-rainy season precipitation. These patterns are interpreted in terms of selection for dispersal and the within-year timing of germination.     

QUANTITATIVE GENETICS OF SIZE,SHAPE, LIFE-HISTORY,AND FRUIT CHARACTERISTICS OF THE SEED-HETEROMORPHIC COMPOSITE HETEROSPERMA PINNATUM. I. VARIATION WITHIN AND AMONG POPULATIONS

We document phenotypic and genetic variation within and among populations of the seed heteromorphic species Heterosperma pinnatum Cav. (Compositae) in the production of seed morphs and in a variety of life-history and morphological characteristics that might be correlated with seed and head traits. Each trait is found to have significant genetic variance in most or, usually, all populations. Significant among-population genetic variation exists for all traits except number of achenes per head and seedling shape, although some traits have much less genetic variation among than within populations. Number and percentage of intermediate achenes per head, total number of achenes per head, and lengths of central and peripheral achenes had little among-population genetic variation compared to within-population variation. Most traits had slightly less genetic variation among than within populations; however, some traits (percentage of central achenes, length of awns, date that the first flowering head opened, date that the first fruiting head opened, and death date) had more among-population genetic variation. The proportions of achene morphs produced had high broad-sense heritabilities and high among-population genetic variance, except in the case of intermediate achenes. All phenological variables had high among-population genetic variation. Within-population heritabilities were high for dates of first flowering head and fruiting head but low for death date and reproductive interval. Family and population means measured in the greenhouse for traits having high broad-sense heritability or among-population genetic variance were closely correlated with field means for the corresponding families or populations. The amounts of phenotypic variation were similar for traits that were measured in both the field and the greenhouse. These lines of evidence suggest that greenhouse results provide reasonable estimates of genetic variation in the field for this species. Numerous studies have reported variation in the proportion of seed morphs for different heteromorphic-seeded species and have discussed adaptive scenarios for the evolution of seed proportions; however, our investigation is one of only a few that have documented the amount of phenotypic and genetic variation within and among populations.     

Dormancy and dispersal in dimorphic achenes of tansy ragwort,Senecio jacobaea L. (Compositae)

Summary Marginal and central florets of the capitula of tansy ragwort Senecio jacobaea yield different kinds of fruit. The central (disk) achenes are lighter (x±SE=199±5g), more numerous (x±SE=58±0.6 achenes per head), and are equipped with a pappus aiding wind transport and rows of trichomes aiding animal transport. The marginal (ray) achenes are heavier (x±SE=286±7g), less numerous (virtually invariant at 13 achenes per head), and lack dispersal structures. Whereas disk achenes are relased shortly after they mature, ray achenes are retained by the parent for a period of months following maturity.Germination at constant temperature (20°C) and with alternating light (12 h light: 12 h dark) demonstrated that disk and ray achenes exhibit different germination syndromes. Germination percentage increases linearly with achene fresh weight in both types; for a given weight, disk achenes have a higher germination percentage than ray achenes. Germination time decreases with increasing achene weight in disk achenes, but increases with achene weight in ray achenes.The germination percentages and germination times for disk and ray achenes diverge progressively with increasing achene weight. The divergence in behavior is a result of diverging patterns of dry matter allocation in the two achene types. Increase in the size of disk achenes favors the embryo fraction, thereby speeding germination while reducing protection. Increase in the size of the ray achenes favors the pericarp fraction, thereby increasing protection while delaying germination.Reduced germination percentage and germination speed of the ray achenes were shown by experimental manipulation to be caused by physical inhibition by their thicker pericarps.Dimorphism in ragwort likely speards germination out in space and time, thereby increasing the number of safe sites an individual parent can exploit in disseminating offspring. The syndrome in other heteromorphic composites resembles that of ragwort, generally combining reduced dispersal-delayed germination in the outer achenes and distance dispersal-quick germination in the central achenes. The outer achenes are generally less numerous and larger. Dispersal traits (large numbers, early release and light wieght) are the direct opposite of dormancy traits (small numbers, delayed release and heavier weight). Thus conflicts between the properties determining dormancy and dispersal appear to require separate dormancy and dispersal phenotypes.     

Inbreeding effects on fitness traits in the heterocarpic herb Leontodon autumnalis L. (Asteraceae)

Heterocarpic plants are characterized by the production of distinct types of fruits that usually differ in their ecological behavior. In the Asteraceae, differences are mainly found between peripheral non-dispersal and central dispersal achenes (single-seeded fruits). Inbreeding depression is considered as an evolutionary force as it may reduce several fitness traits, and in the case of heterocarpic plants, it could influence fitness traits (e.g., seed set, germination rate, growth rate) of each fruit morph, which may have important ecological and evolutionary consequences. In particular, differential effects on fitness traits and dispersal of selfed and outcrossed progeny can strongly determine the viability of extant populations and the potential to colonize new habitats. We conducted a hand-pollination experiment in greenhouse conditions to test whether inbreeding affects the fitness of achene morphs in the heterocarpic herb Leontodon autumnalis (Asteraceae). Results show that achene morphs significantly differ in their ecological behavior, peripheral achenes germinating more and faster than central achenes. The significant interaction between pollination treatment and achene morph for germination probability might indicate a link between dormancy and mating system in L. autumnalis: germination was higher for outcrossed achenes in central achenes whereas the opposite pattern was exhibited by peripheral achenes. Selfing dramatically reduced seed set, probably as a consequence of strong self-incompatibility mechanisms rather than inbreeding effects. Inbreeding depression significantly affected late life-cycle traits, such as growth rate and biomass at flowering. Overall, results suggest that inbreeding depression seems to be an important selective force maintaining outcrossing in L. autumnalis.     

Changes in seed dormancy of Rosa multibracteata Hemsl. & E. H. Wilson with increasing elevation in an arid valley in the eastern Tibetan Plateau

Variations in levels of seed dormancy among conspecific populations are poorly understood. We aim to determine the variations in the level of dormancy in Rosa multibracteata along an elevational gradient, and to analyze the mechanisms underlying seed dormancy. The study was conducted in an arid valley in the eastern Tibetan Plateau at five elevations: 1700, 1900, 2100, 2300 and 2500 m. Achene traits were measured and physiological levels of dormancy were determined by measuring germination percentages of achenes treated with H₂SO₄ scarification, warm stratification and cold stratification and combinations of those treatments. Achene size, mass and pericarp thickness increased with increasing elevation, but embryo width decreased with increasing in the arid valley. The level of dormancy generally increased with increasing elevation. The level of dormancy of the rose achenes is intermediate from sites at elevations of 1700, 1900 and 2100 m and deep at elevations of 2300 and 2500 m. Comprehensive analysis suggests that the elevational pattern of level of seed dormancy could be ascribed to intrinsically thicker pericarp and deeper physiological dormancy in the embryo, and extrinsically decreased temperature, increased precipitation and soil moisture at the higher elevations. The great variations of achene traits and levels of seed dormancy across elevations suggest a phenotypic differentiation exists within this rose species. Thus, habitat conditions and achene traits should be considered when selecting roses as materials used to propagate seedlings for ornamental purposes or plant restoration.     

Variability of the inflorescence ofMicroseris laciniata (Compositae: Cichorieae)

Quantitative characters of the flowering head of a garden population ofMicroseris laciniata were scored during the second, third, and fourth season of growth. Number of achenes per head, number of phyllaries per head and the average number of pappus parts per achene in single heads show significant plant to plant variation. Achenes per head and pappus parts per achene were scored in identical plants in two subsequent seasons. The number of pappus parts per achene varies freely between five and ten. This contrasts with annual species ofMicroseris in which either five or ten pappus parts are found, depending on the species. In spite of a clear plant-specific average of pappus parts, both high and low pappus part determination can be demonstrated in all specimens. The number of pappus parts depends on the position of an achene on the receptacle, marginal achenes usually having fewer pappus parts than central ones. This gradient is not closely correlated with the position of an achene on the genetic spiral.     

一年生短命植物疏齿千里光果实异形性的生态学意义

疏齿千里光(Senecio subdentatus)是分布在新疆北部古尔班通古特沙漠中的一种具异形瘦果的菊科一年生短命植物。将野外观测与室内实验相结合, 对该物种异形瘦果的形态、扩散和萌发特性, 以及异形瘦果产量与植株大小的关系进行了研究, 并对其生态学意义进行了探讨。结果表明: 疏齿千里光果序中的外围果和中央果均为柱形, 但前者为淡黄色, 后者为褐色, 且二者在大小、冠毛长度及果皮微形态等方面均存在明显差异。两种瘦果均以单个果实为扩散单元, 且在静止空气中的降落速度和在1 m·s^-1与2 m·s^-1风速下的扩散距离无显著差异, 说明虽然果实大小和冠毛长度对瘦果扩散具有不同的影响, 但对其整体扩散能力无明显影响。在各温变周期(5/2、15/2、20/10、25/15和30/15 ℃)处理中, 淡黄色外围果的萌发率均高于褐色中央果的, 且不同温度间两种瘦果的萌发率均存在显著差异, 但光照条件对其无显著影响。果序中的中央瘦果数明显多于外围果的, 且植株中外围果所占比例与植株大小间呈显著负相关关系, 而中央果所占比例与植株大小间呈显著正相关关系。这些特点说明, 该物种的小植株倾向于产生较多较易萌发的外围果, 大植株倾向于产生较多不易萌发的中央果。在古尔班通古特沙漠不可预测的极端环境中, 疏齿千里光可通过异形瘦果间的萌发差异及调节其不同大小植株中异形瘦果的比例, 来减少同胞后代之间的竞争, 增加其对不同微环境条件的生态适应性。     

Variability and Cryptic Heteromorphism of Ambrosia artemisiifolia Seeds: What Consequences for its Invasion in France?

BACKGROUND AND AIMS: Ambrosia artemisiifolia is a ruderal weed introduced from North America to Europe. It produces large amount of achenes which are highly heterogeneous in size. Due to the preponderant role of propagules in invasive plant processes, the achene mass variability related to germination, dispersal strategy and life history traits of offspring were investigated within this species. METHODS: The variability in achene mass was quantified among six populations sampled in different habitats. The effects of achene mass variation on germination were studied. The percentages of floating and non-floating achenes were evaluated in the studied populations. The consequences of floatability on the growth and traits of the offspring were studied. KEY RESULTS: Mean achene mass ranged from 1.72 to 3.60 mg, depending on the populations, and was highly variable. Variation among achenes within plants accounted for 63.9 % of the variance, whereas variances among plants within each population (22.2 %) and among populations (13.9 %) were lower. Achene masses were also positively correlated to the total germination percentage for four populations out of six. Two kinds of achenes were distinguished: floating and non-floating. The majority of floating achenes (90 %) sank 24 h after water immersion. Whatever the population, floating achenes were lighter, more dormant and germinated faster than non-floating achenes. Plants which issued from floating achenes had better growth than those from non-floating achenes. CONCLUSIONS: The capacity of A. artemisiifolia to be invasive in Europe appears to be high, possibly due to its huge plasticity in seed mass which may help it to cope with a wide range of conditions and to establish in disturbed habitats. Furthermore, the recent invasion of southern France by A. artemisiifolia could be partially explained by water dispersal of achenes through rivers and has pinpointed its colonization potential along French rivers.     

Testing achene germinationof Potamogeton praelongus Wulfen

The goal of this work was to determine the best method of breaking the achene dormancy in Potamogeton praelongus Wulfen. The ways of breaking achene dormancy studied in this experiment included methods of achene storage, stratification, UVA radiation, anaerobic conditions, mechanical disruption of achenes?outer layers and their chemical disruption by NaClO. Nine different treatments of achenes were combined with two methods of achene storage. Particular achene treatments and storage conditions were proven to have a significant impact on breaking dormancy. Although the highest germination rate (83.3%) was achieved when the dormancy was broken chemically by long effect of 100% concentrations of Savo detergent (containing 5% NaClO), the growth of the sprouts was subsequently inhibited due to toxic effects of Savo. Thus the most successful treatment was based on changing temperature, e.g. 2.5 months of cold storage followed by 14 days at room temperature (germination rate 32.7%). This treatment was also most similar to the natural process. Germinated achenes were also found in Petri dishes exposed to UVA radiation, anaerobic conditions and chemical disruption of the outer layers. Results of these treatments were influenced by the storage method.     

Significance of achene characteristics and within-achene resource allocation in the germination strategy of tetraploid Aster pilosus var. pilosus (Asteraceae)

The interrelationships among achene weight, allocation to embryo and pericarp, and germination time were determined for 500 stratified achenes of tetraploid Aster pilosus Willd. var. pilosus. Only 52.6% of the achenes germinated. Germinated achenes were significantly heavier than ungerminated achenes. Germination time was independent of achene weight and embryo weight, but varied inversely with pericarp weight. Variable achene weight is evolutionarily advantageous. Heavy achenes are at an advantage in that their proportionately larger embryos and thinner pericarps facilitate germination, promoting competitive establishment of seedlings. Lighter achenes are also at an advantage through increased dispersibility, and their relatively thick pericarp provides a persistent seed bank. Evolutionary pressures presumably maintain the variability in achene weight of var. pilosus. These results are discussed in the context of the early midsuccessional ecology of var. pilosus.     

Morphometric Analysis of Sunflower (<Emphasis Type="Italic">Helianthus annuus</Emphasis> L.) Achenes from Mexico and Eastern North America<Superscript>1</Superscript>

Morphometric Analysis of Sunflower ( Helianthus annuus L.) Achenes from Mexico and Eastern North America. Sunflower (Helianthus annuus L.) has played a major role in the evolution of agricultural systems in the Americas. The discovery of ancient domesticated remains from archaeological deposits in pre-Columbian Mexico offers new dimensions to widely accepted viewpoints on the domestication pattern of H. annuus. Although American sunflower populations north of Mexico have been examined extensively, Mexican indigenous domesticated landraces have not been studied in any detail. In this study, we morphologically assessed wild and domesticated sunflower achenes from Mexico and compared them to similar datasets from eastern North America. Additionally, we evaluated the utility of four computer-assisted shape measurements in discriminating between wild and domesticated sunflower achenes (fruits) and compared variation in achene size among modern wild and cultivated populations from both Mexico and the U.S. We found that, of the shape parameters tested, none were informative in distinguishing wild achenes from domesticated varieties. Subsequent size analysis, using conventional parameters of length, width, and thickness, showed that modern wild populations from Mexico had smaller achenes compared to modern populations from eastern North America. Domesticated achenes unearthed from Mexican archaeological sites, however, were significantly larger than the early domesticated specimens recovered from eastern North America. Our methodological results indicate that variation in archaeological sunflower achenes is better described by conventional size parameters rather than computerized shape analysis.     

Ecology of achene dimorphism in Leontodon saxatilis

BACKGROUND [corrected] AND AIMS: Leontodon saxatilis produces two morphologically distinct achenes (morphs) in a single capitulum: one row of dark brown achenes without a pappus lies at the edge ('peripheral achenes'; 0.74 +/- 0.18 mg) while the inner ones are light brown with a pappus ('central achenes'; 0.38 +/- 0.07 mg). The hypothesis that achene heteromorphism in L. saxatilis widens its ecological amplitude was tested. KEY RESULTS: Achenes of both morphs germinated over the same range of temperatures (6-33 degrees C) but the central achenes showed significantly higher germination percentages, and the two also differed significantly in their annual dormancy cycle, with the peripheral achenes showing greater dormancy for part of the year. Seedlings from the two morphs did not differ significantly in total biomass after 2 and 4 weeks of growth, neither did they differ significantly in root and shoot weight and root:shoot ratio. Plants from both morphs growing at different regimes of soil moisture, nutrients and competition did not differ significantly in their number of achenes per capitulum. While the number of central achenes varied, that of peripheral achenes remained constant at approx. 13. Drier soil led to an increase in the number of central achenes in plants from both morphs. CONCLUSIONS: The peripheral achenes can replace the mother plant in the following growing season, whereas the central achenes are well adapted for wind dispersal and thus for colonization of new sites. However, the extent of variation in germination characteristics was similar to that found in seed populations of homomorphic plants, which suggests that germination percentage and other growth characteristics do not contribute to widening the ecological amplitude. The habitat of most heteromorphic species, the morphs of which differ greatly in germination and/or growth characteristic, are deserts or highly disturbed areas where such differences are highly advantageous, but in the moderate habitat of L. saxatilis the differences may prove a disadvantage.     

Ecological correlates of achene mass variation in Chrysothamnus nauseosus (Asteraceae)

Chrysothamnus nauseosus (Asteraceae) is a complex polymorphic shrub species widely distributed in western North America. In a study of 86 populations belonging to 15 subspecies, achene mass varied among populations over a sevenfold range. Achene mass was closely correlated with capitulum size at the subspecies level, varied little between wild-growing and common-garden-grown members of a population, and was under strong genetic control. Subspecies with the heaviest achenes are restricted to specialized edaphic environments (dunes and badlands) or late seral montane riparian communities, while subspecies that are widely distributed and that occur in early seral habitats have less heavy achenes. Selection on achene mass has apparently been a notable feature of the adaptive radiation of Chrysothamnus nauseosus into the wide array of habitats it currently occupies. Within a wild population, achene mass was greater for plants fruiting in midautumn than for plants fruiting early or late in the autumn, and this same trend was observed within individual plants in garden populations, indicating environmental control, perhaps through resource limitation. Highly significant between-plant differences in achene mass were found in both wild and garden populations, suggesting that within-population genetic variation is sufficient to permit continuing selection.     

中国特有植物艾比湖沙拐枣(Calligonum ebi-nuricum)居群内果实性状的变异

以中国荒漠特有植物艾比湖沙拐枣(Calligonum ebi-nuricum)果实形态特征为研究对象,对分布于原产区的同一居群12个样株的果实进行形态特征比较研究。通过每株30粒果实的喙长、果实长度、果实宽度、果实形状、瘦果长度、瘦果宽度、瘦果形状、刺毛长度、头部刺毛长度、两肋间距、两刺间距11个性状指标的数值分析表明：在株内与株间均有变异,株间果实性状变异较株内果实性状变异大;代表果实和瘦果大小指标正相关性极显著（p＜0.01）;两肋间距与瘦果形状呈负相关（r=-0.297**,p＜0.01）。研究结果证明：其果实形状和瘦果形状（长度、宽度及长宽比）较为稳定,可作为艾比湖沙拐枣的分种依据,而其它果实性状可塑性较大,只能作为分类的参考。该研究结果可供沙拐枣属其它种的准确分类参考。     

Morphological variability of achenes of some European populations of Lactuca serriola L.

This study focuses on the morphological variability of Lactuca serriola achenes in relation to eco-geographic features. Fifty L. serriola populations from four European countries, Czech Republic, Germany, the Netherlands and United Kingdom, were studied. Five morphological characters of the achenes - length and width of achene, length/width index, length of beak, and number of ribs - were evaluated. Significant differences exist in achene morphology of two leaf forms of L. serriola, forma serriola and forma integrifolia. Achenes of f. serriola are shorter, thinner, shorter beaked, lower length/width index, and higher number of ribs compared to f. integrifolia. There was significant variation in the measured characters. Statistical analysis indicated that achene length and width increased along an east-west transect from 2.95 to 3.35 mm and 0.93 to 1.00 mm, respectively. Mean beak length had a similar trend with the exception of German achenes. They had shorter beaks than achenes originating from the Czech Republic with 4.38 and Germany with 4.33 mm. The same trend was evident for L/W index from Czech with a ratio of 3.21 and Germany with 3.14. The number of ribs increased from east to the west in continental Europe, whereas the lowest number of ribs was recorded in achenes collected in Czech with 10.89 and the UK with 10.59. Achene morphology was significantly correlated with three eco-geographic features; longitude, latitude, soil texture of the habitats. The other eco-geographic factors, altitude and population size, did not significantly correlate with the studied characters of L. serriola achenes.     

Germination requirements and dispersal timing in two heterocarpic weedy asteraceae

In SW Spain the winter annuals Anacyclus radiatus and Chrysanthemum coronarium (Asteraceae) are found as weeds in diverse crops. Both plant species are heterocarpic, i.e. the peripheral and central achenes of the capitulum are morphologically distinctive. In heterocarpic and heterospermic species the different fruit or seed morphs usually have differential ecological behaviour. In this work we have studied the morphometry, germination and dispersal timing of t he different achene morphs in A. radiatus and C. coronarium. Laboratory germination tests were carried out to evaluate the influence of incubation temperature (light/dark, 27/27, 24/18, 20/10, 15/5, 10/4 degrees C), chilling period (0, 1, 7, 21 and 60 days at 2 degrees C), GA3, nitrate and the dark on the germination percentage and rate (t50). The peripheral achenes of A. radiatus have a significantly broader wing than the central achenes. In this species, germination was inhibited in the dark but viable achenes of both types germinated completely under light exposure irrespective of test conditions. Nevertheless, the peripheral achenes germinated significantly faster compared to the central achenes (t5o 1.04 vs. 1.55 days at 24/18 degrees C). In C. coronarium peripheral achenes have three-winged ribs and are significantly longer, wider and thicker than the central achenes, which have only one- or two-winged ribs. In this species the peripheral achenes showed a much lower germination compared to the central achenes under all conditions tested (0.3-3.5% vs. 41.5-58.0%). Embryos isolated from the pericarp of peripheral achenes germinated quickly in a great extent, and when placed in close contact with a pericarp fragment they did not show any inhibition of germination, suggesting physical dormancy. To determine differences in dispersal timing between achene morphs, we monitored dispersal in a stand of both plant species. In A. radiatus the peripheral achenes were shed first, at the onset of the rainy season in late summer, whereas the central achenes dispersed from late summer to early winter. In contrast, in C. coronarium the central achenes were shed first, from early summer to early autumn, whereas the peripheral achenes dispersed mainly in early autumn. Thus, in both weed species the more dispersible and readier-to-germinate achene morph is dispersed first. The differences in the germination and dispersal behaviour between achene morphs represent an opportunistic, mixed strategy which is favourable under environmental uncertainty, and should be accounted for in order to understand the population dynamics of both weed species.     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

Effects of pollen donors on seed formation inEspeletia schultzii (Compositae) populations at different altitudes

The influence of different pollen donors on seed formation was investigated in three populations ofEspeletia schultzii that differ in environmental conditions and life history characteristics. Self pollen and pollen from different donors (< 15m apart) within each population was used in a diallel design in order to test the genetic base of seed set variation. Three measures of seed formation were used: (1) achene number; (2) proportion of filled achenes (fruits) that distinguishes between achenes with seeds and empty achenes; (3) proportion of aborted seeds that distinguishes between viable and aborted seeds. Self-pollinations resulted in empty achenes. Achene number did not vary between the different pollen donors. A bimodal pattern of filled achenes was found in two populations in two consecutive years. On the other hand, a unimodal pattern was found in crosses between more distant donors (> 30m). These patterns seems to be the results of a sporophytic incompatibility system. Seed abortion was highest at the higher elevations and seems to be correlated with elevation rather than with any genetic effect.     

DIFFERENTIAL GERMINATION OF RAY AND DISC ACHENES IN HEMIZONIA INCRESCENS (ASTERACEAE)

The differential germination responses of ray and disc achenes of Hemizonia increscens (Asteraceae) were compared in field and laboratory investigations in order to gain insight into the ecological and evolutionary significance of heterocarpy. In the field, 200 ray and 200 disc achenes were placed in native, sterilized soil in a series of cleared, randomized, replicated plots. In a nearby plot a similar number of achenes were placed in plastic petri dishes in which high moisture conditions were maintained. Disc achene germination occurred under relatively minimal moisture conditions (<1 cm rainfall for 19 days) and relatively mild temperature regimes (21–7 C). Disc achene germination began three days after planting in the field plot and four days after they were put into the field petri dishes. In contrast, the onset of ray achene germination occurred 21 days after planting in the field plot and 19 days after planting in the field petri dishes. Averages of 2.05 and 2.71 disc achenes/day germinated in the field plot and field petri dishes, respectively. These contrasted with averages of 0.57 and 0.50 ray achenes/day germinated in the field plot and in the field petri dishes, respectively. A total of 67.5% and 69% disc achenes germinated in the field plot and the field petri dishes, but only 18% and 16.5% ray achenes germinated in the field plot and field petri dishes, respectively. Three separate treatments, using 100 ray and 100 disc achenes in each, were performed in laboratory growth chambers: 1) nicking the fruit coat, 2) excising the embryo, and 3) leaving the fruit coat intact. Onset of germination for all disc achene treatments occurred after three days. No significant differences were found among the three disc achene treatments in timing, rate, or germination percentage. All three disc treatments in the laboratory closely paralleled those for disc achenes in the field plots in time and germination percentage, but rates of germination were not as high. Germination of the nicked and excised ray achenes treatments began after four days, while germination of the untreated ray achenes began after 27 days. Untreated ray achenes in the laboratory paralleled the ray achenes in both field experiments in rate and germination percentage, but were delayed in time of germination. The nicked and excised ray achene treatments, however, were similar to the disc achene treatments in time of germination, and were not significantly different from disc achenes in rate of germination. These data suggest 1) that ray and disc achenes are markedly different in germination under identical conditions in field and laboratory experiments, and 2) this difference in germination response may be due to the thicker pericarp of the ray achenes.     

山东莎草属植物果实形状及果皮微形态研究

利用扫描电镜对山东分布的18种莎草属植物的果实形状及果皮微形态进行了观察研究。结果表明,莎草属植物果实为椭圆状、倒卵状或卵状三棱形,顶端具柱状或锥状短喙;果皮微形态可分为:网-疣复合纹饰、网状纹饰、疣状纹饰和无规则突起纹饰4种类型;果皮微形态类型的划分与基于宏观形态特征的传统分类关联较小,但在该属形态相近的种间存在明显差异,可作为种间划分的依据。果皮微形态证据支持将水莎草属(Juncellus)、砖子苗属(Mariscus)和断节莎属(Torulinium)归入莎草属(Cyperus)的分类观点。