Proximate Limitations to Fruit and Seed Set in Phellodendron amurense var. sachalinense

Abstract Fruit and seed set in a dioecious tree, Phellodendron amurense var. sachalinense grown in isolation, were studied for three years in Hokkaido, northern Japan. Hand pollination did not affect fruit set during the three years, but did affect seed-set by increasing the proportion of fruits with higher seed numbers. Artificial defoliation resulted in decreased mean seed mass, but caused no significant differences in mean seed number per fruit. Total seed mass per infructescence increased with inflorescence size except with artificial defoliation where it reached an upper limit. Although the results of the artificial defoliation treatment indicate that resources are most important for fruit and seed set, it is concluded that fruit and seed set of the tree are not limited by resources, since the number of fruits increased with inflorescence size and an artificial defoliation treatment did not decrease seed-number.     

RE-EVALUATING THE SIGNIFICANCE OF CORRELATIONS BETWEEN SEED NUMBER AND SIZE: EVIDENCE FROM A NATURAL POPULATION OF THE LILY,CLINTONIA BOREALIS

The basis for the negative correlation between seed number and seed size was experimentally investigated in a natural population of Clintonia borealis. Clones of this species vary significantly in estimated self-compatibility (ratio of seed set with selfing to that with outcrossing) and this appears to affect the number and size of seeds set in individual flowers of each. Clones estimated to be largely self-compatible set more seeds per flower than incompatible ones under natural pollination. However, naturally pollinated flowers of self-compatible clones set smaller seeds than those of incompatible clones, and the significance of the negative relationship between seed number and size in individual flowers was removed by holding variation due to compatibility constant. Supplementing resources per flower (by reducing the number of fruits competing for resources per stem) significantly increased total seed mass but had no effect on the negative relationship between seed number per flower and seed size. In contrast, supplementing cross pollination did not significantly influence total seed mass per flower but changed the relationship between seed number and size to positive, regardless of resource level. In other words, with plentiful cross pollination maternal genets capable of setting more seeds per flower also produced heavier ones. Thus, evidence is provided that the balance between seed number and seed size in this population is regulated by the interaction of maternal self-compatibility with natural pollination.     

Cytokinins in the perianth, carpels, and developing fruit of Helleborus niger L

Reproductive development in the Christmas rose (Helleborus niger L.) differs from that in commonly investigated model plants in two important aspects: (i) the perianth develops a photosynthetic system, after fertilization, and persists until seed ripening; and (ii) the ripe seed contains an immature embryo which continues to mature off the mother plant. The possible roles of cytokinins in these processes are investigated here by analysing extracts of the perianth and the carpels/maturing fruit prepared during anthesis and four stages of post-floral development. trans-Zeatin, dihydrozeatin, N6-(Delta2-isopentenyl)adenine, and their ribosides were identified by tandem mass spectrometry. Single ion monitoring in the presence of deuterated internal standards demonstrated the additional presence of the corresponding riboside-5'-monophosphates, O-glucosides, and 9-glucosides, and afforded quantitative data on the whole set of endogenous cytokinins. Fruit cytokinins were mostly localized in the seeds. Their overall concentrations increased dramatically during early seed development and remained high for 6-8 weeks, until shortly before seed ripening (the last time point covered in this work). Overall cytokinin levels in the perianth did not change markedly in the period covered, but the level of N6-(Delta2-isopentenyl)adenine-type cytokinins appeared to increase slightly and transiently during the greening phase. The perianths of unpollinated or depistillated flowers, which survived, but did not pass through the complete greening process, contained significantly less cytokinins than observed in fruit-bearing flowers. This suggests that perianth greening requires defined cytokinin levels and supports the role of the developing fruit in their maintenance.     

Effects of parentage,prior fruit set and pollen load on fruit and seed production in Campanula americana L

Summary We conducted a controlled crossing experiment to examine the effects of maternal and paternal parentage, the size of the pollen load, and prior fruit production on the proportion of flowers that set fruit, seed number per fruit and seed weight in a natural population of Campanula americana. Effects due to the maternal parent were large for all measures of fruit and seed production, while the paternal parent had a significant effect only upon mean seed weight. As the number of prior fruits on the maternal plant increased the probability that a flower would produce a mature fruit, the number of seeds per fruit, and total seed weight per fruit all decreased. We found no effect of the size of the pollen loads used in this study on fruit or seed production. These results are consistent with those of other studies that suggest in natural plant populations maternal effects, especially environmental maternal effects, can have an overwhelming effect on fruit and seed production and on seed characteristics.     

Plasticity of reproductive traits responding to variation in light availability at the rosette stage of the first year in a strict biennial,Pedicularis torta,from a field on the Qinghai–Tibet Plateau,China

Light is a fundamental heterogeneous environmental factor for plants and often varies at both spatial and temporal scales. In the present study, we carried out a shading experiment to investigate the effect of variable light availability at the rosette stage of the first year in the strict biennial species Pedicularis torta (an endemic species of the Qinghai‐Tibet Plateau) on reproductive traits (i.e. total number of flowers per plant, total number of fruits per plant/total number of flowers per plant, the number of seeds per fruit, mean individual seed mass and estimation of the total number of seeds per plant, plant size at maturity in the second year and on the relationships between reproductive traits and plant size at this stage). We found that light availability affected most of the reproductive traits (except for mean individual seed mass), plant size at maturity in the second year, and the relationship between the total number of flowers per plant and plant size at this stage. Our results indicate that reproductive traits respond to changes in light availability during the rosette plant stage.     

Seed size selection by olive baboons

Seed size is an important plant fitness trait that can influence several steps between fruiting and the establishment of a plant’s offspring. Seed size varies considerably within many plant species, yet the relevance of the trait for intra-specific fruit choice by primates has received little attention. Primates may select certain seed sizes within a species for a number of reasons, e.g. to decrease indigestible seed load or increase pulp intake per fruit. Olive baboons (Papio anubis, Cercopithecidae) are known to select seed size in unripe and mature pods of Parkia biglobosa (Mimosaceae) differentially, so that pods with small seeds, and an intermediate seed number, contribute most to dispersal by baboons. We tested whether olive baboons likewise select for smaller ripe seeds within each of nine additional fruit species whose fruit pulp baboons commonly consume, and for larger seeds in one species in which baboons feed on the seeds. Species differed in fruit type and seed number per fruit. For five of these species, baboons dispersed seeds that were significantly smaller than seeds extracted manually from randomly collected fresh fruits. In contrast, for three species, baboons swallowed seeds that were significantly longer and/or wider than seeds from fresh fruits. In two species, sizes of ingested seeds and seeds from fresh fruits did not differ significantly. Baboons frequently spat out seeds of Drypetes floribunda (Euphorbiaceae) but not those of other plant species having seeds of equal size. Oral processing of D. floribunda seeds depended on seed size: seeds that were spat out were significantly larger and swallowed seeds smaller, than seeds from randomly collected fresh fruits. We argue that seed size selection in baboons is influenced, among other traits, by the amount of pulp rewarded per fruit relative to seed load, which is likely to vary with fruit and seed shape.     

Changes in Baboon Feeding Behavior: Maturity-dependent Fruit and Seed Size Selection within a Food Plant Species

Despite considerable inter- and intraindividual variation in fruit and seed size in many plant species, researchers have given little attention to the relevance of the traits for primate fruit choice within a food plant species and its implications for tree regeneration. We studied feeding behavior and selectivity of olive baboons (Papio anubis) in the African locust bean (Parkia biglobosa, Mimosaceae), via direct observations of habituated groups and indirect evidence from leftovers of pods after feeding events. Olive baboons acted as both seed predators and dispersers for Parkia biglobosa. They fed on and destroyed unripe seeds, and swallowed intact ripe seeds when consuming mature fruit pulp. Predation rate was high, and only 10% of the seeds were dispersed. Predation and dispersal of seeds is linked to seed number and size. Digestible unripe seeds accounted for 10% of the unripe fruit mass, while indigestible ripe seeds made up 28% of the mature fruit mass. With these constraints, olive baboons increased food gain per fruit by selecting unripe pods containing a high number of large and heavy seeds. Consequently, only pods with fewer and smaller seeds remained for maturation. Thereafter, baboons fed on mature pods containing the smallest seeds, and exploited pods with more seeds to a greater extent than those with fewer seeds. Thus, fruits with small seeds and an intermediate seed number contributed the most to dispersal by baboons.     

From flowering to dispersal: reproductive ecology of an endemic plant,Astragalus australis var. olympicus (Fabaceae)

Astragalus australis var. olympicus is an endemic plant of the Olympic Mountains, Washington. It is considered a Species of Concern by the U.S. Fish and Wildlife Service. This study focused on the reproductive biology of the plant from flower production through seed germination to identify possible weak points that might contribute to its rarity and impede its conservation. Most plants produced a large mean number of flowers and ovules (314.8 and 4106, respectively), but relatively few of these formed fruits and seeds (25.8 and 3.8%, respectively). In decreasing importance, ovules in fruits were lost to predation, seed abortion, and lack of fertilization. The percentages of these fates differed among sites and years. Excluding pollinators by bagging flowers reduced fruit set by ∼50%, but seed set per fruit and seed mass were unaffected. Germination was affected by scarification, temperature, and moisture availability. About 11% of seeds damaged by predispersal seed predators (weevil larvae) remained viable and were released from dormancy. I hypothesize that predispersal seed predation (over 80% at one site) has a negative effect on population growth. Conservation of this species could benefit from improved fruit set and decreased seed predation.     

Floral costs in Nigella sativa (Ranunculaceae): compensatory responses to perianth removal

Because internal resources are finite, it has been assumed that attractive, floral organs represent a significant drain on the energy and nutrient budget of a plant. Despite the broad significance of such trade-offs, in relatively few studies have investigators manipulated floral investments, then evaluated allocation to subsequently produced flowers, fruits, and seeds. In the present study of Nigella sativa, the cost of maturing and/or maintaining perianths was documented after all sepals and nectaries were removed at the bud stage and a significant increase in mean seed mass, the total amount of biomass allocated to seed production, and mean germination rate of the maternal seed crop were measured. The increased biomass, carbon, and nitrogen allocated to seeds were similar in magnitude to the reduction in biomass, carbon, and nitrogen invested in sepals and nectaries after perianth removal. Perianth removal did not significantly affect flower production, maternal fecundity, or progeny seed number. Taken together, these observations indicate the potential for selection-mediated through resource trade-offs with seed mass and time to germination-to cause, or at least facilitate, evolutionary reductions in flower size.     

FERTILIZATION DYNAMICS AND PARENTAL EFFECTS UPON FRUIT DEVELOPMENT IN RAPHANUS RAPHANISTRUM: CONSEQUENCES FOR SEED SIZE VARIATION

Seed weight varies significantly within and among fruits of wild radish (Raphanus raphanistrum). To determine sources of this variation, we studied fertilization and seed development following controlled pollinations. Within fruits, central ovules were fertilized prior to distal ovules and attained greater seed size. Ninety-seven percent of the variation in mean seed wt per fruit was explained by an analysis of variance incorporating parental effects, pollination date, and the number of seeds per fruit. We document strong maternal effects on the number of ovules per ovary, the number of fertilized ovules per ovary, the number of seeds per fruit, and mean individual seed wt per fruit. Across females, pollen donor had a slight but significant effect on seed wt; no paternal effects on fertilization rate, zygote number, or seed number per fruit were detected. Within females, with one exception, pollen donor had no significant effect on these components of seed development. Stronger maternal main effects may be due to donor x recipient interactions, cytoplasmic factors, the genetic inequity within triploid endosperm, and/or strict maternal control over resource allocation. The large maternal effects relative to paternal effects may limit the rate at which natural selection acts on paternal traits expressed prior to seed maturation.     

Long-term variability in seed size and seedling establishment of <Emphasis Type="Italic">Maianthemum bifolium</Emphasis>

Relatively few studies conducted in natural plants populations focus on the relationship between seed size and their germination ability and seedling establishment. Maianthemum bifolium is a perennial herb that spreads vegetatively through rhizomatous growth and reproduces through seeds. However, this species is characterized as seed and microsite limited, and under undisturbed conditions seedlings are not noted. The studies were conducted in two populations of M. bifolium in six subsequent seasons. The mean seed mass was negatively correlated both per ramet as well as in the fruit with the number of seeds, and positively with its height and the number of flowers. The long-term mean annual production of seeds in the populations was 37 and 56 seeds per m². The seeds from both populations had similarly high germination abilities that were approximately 90% under laboratory conditions, 60% in garden, and 55% in the natural habitat. Seeds from four size classes were sown and a positive correlation was noted between seedling establishment and the mass of the seeds from which they grew (r_S = 0.64). Seedling survival was also significantly correlated with seed mass.     

Endogenous Gibberellin Profile During Christmas Rose (Helleborus niger L.) Flower and Fruit Development

Gibberellins (GAs) were identified and quantified during flower and fruit development in the Christmas rose (Helleborus niger L.), a native of southeastern Europe with a long international horticultural tradition. Physiologically, the plant differs from popular model species in two major respects: (1) following anthesis, the initially white or rose perianth (formed in this species by the sepals) turns green and persists until fruit ripening, and (2) the seed is shed with an immature embryo, a miniature endosperm, and a prominent perisperm as the main storage tissue. GA₁ and GA₄ were identified by full-scan mass spectra as the major bioactive GAs in sepals and fruit. LC-MS/MS system in accord with previously verified protocols also afforded analytical data on 12 precursors and metabolites of GAs. In the fruit, GA₄ peaked during rapid pericarp growth and embryo development and GA₁ peaked during the subsequent period of rapid nutrient accumulation in the seeds and continued pericarp enlargement. In the sepals, the flux through the GA biosynthetic pathway was highest prior to the light green stage when the photosynthetic system was induced. Unfertilized, depistillated, and deseeded flowers became less green than the seed-bearing controls; chlorophyll accumulation could be restored by applying GA₁, GA₄, and, less efficiently, GA₃ to the deseeded fruit. The sepals of unfertilized and depistillated flowers indeed contained very low levels of GA₄ and gradually decreasing levels of GA₁. However, the concentrations of their precursors and metabolites were less affected. These data suggest that a signal(s) from the fruit stimulates GA biosynthesis in the sepals resulting in greening. The fruit-derived GAs appear to be mainly involved in pericarp growth and seed development.     

Effect of pollen load size and source (self, outcross) on seed and fruit production in highbush blueberry cv. 'Bluecrop' (VACCINIUM CORYMBOSUM; Ericaceae)

Reproductive fitness of a plant is ultimately determined by both number and quality of seed offspring. This is determined by sexual selection of pollen microspores and ovules during pollination and fertilization. These processes may include pollen competition and seed abortion, which reduce the number of microspores and ovules available for final seed production. Thus, even an excess of pollen microspores to ovules does not result in fertile seeds equal to ovule number. We investigated pollen requirements of highbush blueberry (Vaccinium corymbosum cultivar 'Bluecrop') for maximal seed production and how fertile seed number translates into fruit quality, since fruit quality would ultimately determine the dispersal of its offspring. We demonstrate that individual blueberry flowers with a mean of 106 ovules reach their maximum fruit set and mass and minimum time to ripen when 125 outcross pollen tetrads pollinate a flower, compared to 10 or 25. Three hundred tetrads resulted in the increase of fertile seeds, but did not result in a further increase of fruit mass or fruit set, or decrease in time to ripen. We also examined the effect of pure and mixed loads of self and outcross pollen (25 and 125 tetrads), and found no differences in fertile seed number, fruit mass, or percentage fruit set when pollen loads were either 25 self or outcross pollen tetrads, although number of days to ripen was significantly shorter by 8 d with 25 outcross tetrads. When the pollen load of 125 tetrads consisted of self or a 50:50 mixture of self and outcross pollen, fruit mass, days to ripen, and percentage fruit set were not different from loads of 125 outcross pollen. In addition, a pollen load of 25 outcross tetrads resulted in fertile seed number and fruit quality in between that of 25 self, and 125 self, 125 mixed, or 125 outcross tetrads. Large, small, and flat seed types were identified, and only large seeds (length = 1.7 mm) were fertile. These results improve our understanding of pollen load size and source requirements of a crop plant and the limits to pollen transfer when translated to fruit growth.     

Consequences of differences in flowering date on seed production in Heloniopsis orientalis (Liliaceae)

We examined the consequences of differences in flowering date on seed production in the self-compatible herb Heloniopsis orientalis. The number of selfed seeds per fruit, as determined by microsatellite markers, did not depend on when the plant flowered, whereas the number of outcrossed seeds per fruit increased with later flowering dates. Consequently, the selfing rate decreased with later flowering dates. The number of seeds (including both selfed and outcrossed ones) per fruit and the seed?:?ovule ratio increased with later flowering dates. We also examined the effects of pollinators and plant size on seed production. The visitation rate of Diptera did not depend on the flowering season, whereas that of Hymenoptera markedly increased as the flowering season progressed. Diptera stayed longer than Hymenoptera on each plant and flower. Seed production per fruit did not depend on plant size. Thus, the change in selfing rate associated with later flowering dates resulted from the seasonal change in pollinators rather than plant size.     

Long-term variation in seed mass and seed production in populations of Paris quadrifolia

Seed production of the perennial herb Paris quadrifolia L. (Liliaceae) was investigated in five populations in northern Poland. The long-term seed production per square metre differed significantly among populations and years. Moreover, throughout the 7-year study period, 30% of both whole ripe fruits and seeds alone were predated. Variation in seed mass per fruit in both space and time was significant. Throughout the 7-year study, nearly all the marked individuals produced fruits once every 2 years. The most frequent break between fruiting was 2 years and the longest was 5 years. Only the mean seed mass in fruits of the same individual varied significantly over subsequent years. In the five populations, the number of ovules, number of seeds in the fruit and seed mass varied significantly between populations. However seed/ovule ratio did not differ in fruits in the five populations. The seed mass/number trade-off in fruits was strongly partially correlated when the effect of total seed mass was considered. Breeding experiments suggest that P. quadrifolia has a substantial capacity for both inbreeding and outbreeding. There were no significant differences in the seed/ovule ratio, seed number or seed mass in fruits produced from bagged or control flowers. However, both seed/ovule ratio and number of seeds were significantly lower in fruits from emasculated flowers.     

海拔对青藏高原东部毛茛科植物繁殖特征和资源分配的影响

通过野外采样对青藏高原东部高寒草甸上两个海拔间3种常见毛茛科植物条裂银莲花(Anemone trullifo-liavar.linearis)、粗距翠雀花(Delphinium pachycentrumHemsl.)和钝裂银莲花(Anemone obtusiloba)的繁殖性状和资源分配进行了研究,并对植物在极端环境下采取的繁殖策略及繁殖模式进行了探讨.结果显示:(1)3个物种的株高都随着海拔的升高而降低;同一物种的花大小在不同海拔间均无显著差异;条裂银莲花单个种子重随海拔的升高而增加,但虫食数/株、结籽率和种子数/株均随着海拔的升高而减少;钝裂银莲花的繁殖分配、虫食数/株、种子数/株和单个种子重均随着海拔的升高而减小.(2)不同海拔条裂银莲花的个体大小与单花重、雄蕊重、虫食数/株均呈显著正相关,且回归斜率在海拔间有显著差异.粗距翠雀花花期的雄蕊重与个体大小呈显著正相关,但这种异速关系不受海拔和个体大小的影响;钝裂银莲花的个体大小与花期的所有繁殖特征以及果期的种子重/果实均呈显著正相关,不同海拔间个体大小与种子重/果实的斜率差异显著,且与花期各繁殖性状异速关系的截距差异显著.研究表明海拔对植物的个体大小以及种群间的繁殖对策和繁殖成功率有着重要的影响.     

文冠果花粉供体的多样性对果实产出的影响

很多植物的花,其柱头上接受的花粉供体多样性数量通常与果实成熟和种子产出有关。关于提高花粉多样性能否增加植物的生殖产出问题,文献中时常有相互矛盾的结论。本研究通过对自交不亲和性植物文冠果(Xanthoceras sorbifolium)进行1、2、3、4种花粉供体传粉实验,调查了不同花粉供体数量对文冠果雌性生殖的影响。结果表明,增加亲和花粉供体的多样性,一定程度上提高了花序上果实的坐果数量,同时也增加了果实中种子的数量及单粒种子重量。对文冠果授以4种亲和性花粉供体后,花序上成熟果实的平均数最多;3种亲和性花粉供体传粉后,每个果实中的种子数最多,单粒种子也最重。如果混合花粉中含有自交花粉,那么果实将败育。除花粉多样性影响雌性生殖产出外,母本基因型也具有重要影响,即增加文冠果种植品系的多样性,将有助于提高果实和种子产量。本研究还讨论了花粉遗传多样性效应、母本交配选择及果实选择性败育假说。     

TEMPORAL VARIATION OF FLOWER AND FRUIT SIZE IN RELATION TO SEED YIELD IN CELANDINE POPPY (CHELIDONIUM MAJUS; PAPAVERACEAE)

Ecological and evolutionary studies typically consider variation in single reproductive characters in isolation, without considering how they might be correlated with other reproductive and vegetative characters. In our study, we examined temporal patterns of variation and correlation in flower diameter and fruit length during a reproductive phase in two Massachusetts populations of the herb, Chelidonium majus. We also examined the relationships of such variation to measurements of seed yield components (mean seed weight and number per fruit) and aspects of plant vegetative size. Most of the variation in the sizes of reproductive characters occurred within individual plants, instead of among plants or between populations. Flower and fruit sizes as well as seed number per fruit declined significantly during the season in both populations. Only mean seed size per fruit was relatively stable for individual plants in both populations. Conserving resources by a gradual reduction in the size of reproductive characters over the season may be a strategy for maternal plants to continue seed production. The strong, persistent patterns of correlation between certain characters, such as flower and fruit size, in spite of extensive phenotypic plasticity, was interpreted as indirect evidence for developmental correlation. Furthermore, vegetatively larger plants produced not only more flowers and fruits, but also consistently larger flowers and fruits. The results emphasize that variation in fitness characters, such as seed size and number, should not be viewed in isolation from vegetative characters, flower, and fruit sizes in ecological and evolutionary studies, if the goal is to understand the mechanisms of natural selection in wild populations.     

Are large frugivorous birds better seed dispersers than medium‐ and small‐sized ones? Effect of body mass on seed dispersal effectiveness

Frugivorous birds vary in seed dispersal effectiveness (SDE) depending on their body mass. It has been suggested that large birds are more effective dispersers than small ones because they consume a large number of fruits, disperse seeds of distinct sizes, and transport seeds over long distances. Yet, few studies have evaluated the impact of body mass on SDE of birds. In this study, we compiled one database for the quantity (i.e., frequency of visits to plants and number of seeds removed per visit) and quality (i.e., germination of seeds after gut passage and gut retention time of seeds) of seed dispersal by frugivorous birds to evaluate the impact of body mass on SDE. In addition, we compiled data on plant characteristics such as life‐form, fruit type, number of seeds per fruit, and size of seed to evaluate their influence on the quantity and quality of seed dispersal. Data were analyzed with linear mixed effects models and quantile regressions to evaluate the relationship between body mass of birds and quantity, quality, and SDE, in addition to the influence of plant characteristics on SDE. The body mass of birds was negatively related to the frequency of visits to plants. Furthermore, it was positively related to the number of seeds removed per visit, although negatively related to seed size. The life‐form of plants was the only factor explaining the germination of seeds after gut passage. Yet, the body mass of birds was positively related to the gut retention time of seeds. Small and medium birds have a relatively higher SDE than large birds. These results differ from the assertion that large birds are more effective dispersers of plants. Small and medium birds are also effective dispersers of plants that should be preserved and protected from the impact of human activities.