Pollen viability assessments in blackberries (Rubus subgen.Rubus)

Pollen viability has been investigated in 20 blackberry species using 3 methods, (1) cotton blue, (2) TTC, and (3) germination in a sucrose solution. Significant differences were found between species. Correlations between high pollen viability and high seed set were also obtained.     

Apomixis versus sexuality in blackberries (Rubus subgen.Rubus,Rosaceae)

InRubus L. a connection seems to exist between the degree of meiotic disturbances on the one hand, and the production of unreduced embryo sacs, pollen fertility and relative seed set on the other hand. Severe meiotic disturbances commonly encountered in apomictic taxa decrease pollen fertility and thereby seed set since pollen is necessary for endosperm development. By contrast interspecific hybrids between apomictic taxa appear to be sexual and exhibit high pollen fertilities, probably due to an improved meiosis. Thus, apomixis leads to a decreased fertility inRubus, not the opposite, as often discussed.     

Temporal and spatial patterns in the production of pollen, ovules and seeds in pseudogamous blackberries (Rubus subgenus Rubus)

Significant variation in the number of stamens and ovules per flower, the stamen/ovule ratio, and the number of seeds per berry was found among seven blackberry species, each represented by one clone, and to a much lesser extent among inflorescences within species. Inflorescence size and architecture were found to have no major influence on these reproductive parameters.
The position of a flower within the inflorescence strongly influences its onset of anthesis, which in turn is closely correlated with these reproductive parameters. Ovule number decreases with time, although in some species it increases again at the end of the flowering season. Number of seeds decreases more or less linearly with time, although there is an unexpectedly high number of the earliest berries of the triploid species, which is probably the result of interspecific pollination. Pollen production estimated in one species decreases with time, per theca, per flower and per ovule, whereas the percentage of good pollen grains remains unaltered.     

Sources of variation in pollinator contribution within a guild: the effects of plant and pollinator factors

Among plants visited by many pollinator species, the relative contribution of each pollinator to plant reproduction is determined by variation in both pollinator and plant traits. Here we evaluate how pollinator movement among plants, apparent pollen carryover, ovule number, resource limitation of seed set, and pollen output affect variation in contribution of individual pollinator species to seed set in Lithophragma parviflorum (Saxifragaceae), a species visited by a broad spectrum of visitors, including beeflies, bees and a moth species. A previous study demonstrated differences among visitor species in their single-visit pollination efficacy but did not evaluate how differences in visitation patterns and pollen carryover affect pollinator efficacy. Incorporation of differential visitation patterns and pollen carryover effects —commonly cited as potentially important in evaluating pollinator guilds — had minor effects (0–0.6% change) on the estimates of relative contribution based on visit frequency and single-visit efficacy alone. Beeflies visited significantly more flowers per inflorescence than the bees and the moth. Seed set remained virtually constant during the first three visited flowers for beeflies and larger bees, indicating that apparent pollen carryover did not reduce per-visit efficacy of these taxa. In contrast, Greya moth visits showed a decrease in seed set by 55.4% and the smaller bees by 45.4% from first to second flower. The larger carryover effects in smaller bees and Greya were diminished in importance by their small overall contribution to seed set. Three variable plant traits may affect seed set: ovule number, resource limitation on seed maturation, and pollen output. Ovule number per flower declined strongly with later position within inflorescences. Numbers were much higher in first-year greenhouse-grown plants than in field populations, and differences increased during 3 years of study. Mean pollen count by position varied 7-fold among flowers; it paralleled ovule number variation, resulting in a relatively stable pollen:ovule ratio. Resource limitation of seed set increased strongly with later flowering, with seed set in hand-pollinated flowers ranging from 66% in early flowers to 0% in the last two flowers of all plants. Variation in ovule number and resource limitation of seed maturation jointly had a strong effect on the number of seeds per flower. Visitation to early flowers had the potential to cause more seed set than visitation to later flowers. Overall, the most important sources of variation to seed production contribution were differences among pollinators in abundance and absolute efficacy (ovules fertilized on a single visit) and potentially differential phenology among visitor species. These effects are likely to vary among populations and years.     

云南草寇花序内不同部位的性分配

两性花植物花序内的性分配常存在差异,资源竞争、结构效应、交配环境(雌雄异熟、传粉者定向访花行为等)或授粉不均匀等几种假说可以解释这种现象。为验证上述假说,该研究以云南草寇两种表型(雄先熟型和雌先熟型)为材料,分析了其花序内不同部位(基部、中部和顶部)的每花花粉数、胚珠数、花粉/胚珠比、结实率和结籽率,花序内传粉者的定向访花行为,以及人工辅助授粉和去花处理对结实率和结籽率的影响。结果表明:两种表型花序内每花花粉数不随部位而变化,每花胚珠数、结实率和结籽率由基部到顶部依次降低,每花花粉/胚珠比由基部到顶部依次增加,表明顶部花存在偏雄的性分配。人工辅助授粉后,结实率、结籽率仍由基部到顶部依次降低,表明授粉不均匀假说不能解释云南草寇花序内不同部位结实率、结籽率的差异。去除基部和中部花后,顶部花人工辅助授粉条件下的结实率、结籽率与基部花人工辅助授粉条件下的结实率、结籽率无差异,表明云南草寇花序内不同部位结实率、结籽率的差异主要由资源竞争引起。雌先熟表型每花花粉数、花粉/胚珠比高于雄先熟表型,表明两种表型存在性分配差异。传粉者主要先访问云南草寇基部的花,然后向顶部移动。云南草寇花序内顶部偏雄的性分配可能是由资源竞争和传粉者定向访花造成的。     

Cytogenetical and geographical aspects of sterility in Lysimachia nummularia

The probable causes for the high degree of seed sterility often reported for Lysimachia nummularia are discussed. It seems that meiotic irregularities, i.e. lack of bivalent formation, result in the formation of a high percentage of non-viable pollen grains and pollen grains of unequal chromosome number. As demonstrated by the existence of individuals showing seed set, these meiotic irregularities do not lead to total sterility. In a given population, the self-incompatibility of the species and the clonal nature of the population due to its vegetative propagation seem to be a further cause for the sterility observed. The frequency of fertile individuals is comparatively high mainly in Southeast Europe. Probably this area served as a refugium in glacial times. It seems possible that the spread of L. nummularia into northern latitudes resulted in the degeneration of the sexual system. This might have been caused either by the participation of only one or few genotypes spreading mainly vegetatively - in this situation the frequency of seed set would have decreased due to the self-incompatibility of the species - or from the absence of a pollinator, also leading to decreased seed set.     

Is the dichogamy ofSalvia verbenaca (Lamiaceae) an effective barrier to self-fertilization?

Though dichogamy is generally interpreted as a means of preventing self-fertilization, the efficiency of this mechanism has been questioned. Little attention has been paid to functional aspects of male/female timing, such as variation over time in pollen germinability and in stigma receptivity. In the present study these aspects of male/female timing were investigated in the protandrous speciesSalvia verbenaca. To evaluate the effects of selfing, fruit set, total seed weight produced and seed/ovule ratio were also determined, (a) following artificial crossing, (b) following selfing (i.e. bagging) and (c) under natural conditions. Both pollen germinability and stigma receptivity peaked on the last day (day 3) of the flower's life, which presumably favours selfing. Neither fruit set nor seed/ovule ratio were significantly affected by pollination regime, but the total seed weight produced by artificially crossed plants was significantly higher than that produced by bagged or untreated plants. Neither fruit set, seed/ovule ratio nor total seed weight differed significantly between the two years of study. Thus, if aS. verbenaca flower receives outcrossing pollen it will produce a greater weight of seeds (with consequent advantages in terms of female reproductive success). However, the peaking of both pollen germinability and stigma receptivity on day 3 of the flower's life means that selfing is likely to occur if outcrossing pollen has not been received over the early part of the flower's life.     

Pollination intensity and potential seed set in Passiflora vitifolia

Summary Initial seed set and fruit set were pollen-limited in a Costa Rican population of Passiflora vitifolia, a self-incompatible species with 200–350 ovules per flower. Pollination intensity was measured by counting the number of allogamous pollen grains on stigmas of the large one-day flowers. Hand-pollinations demonstrated that 25–50 pollen grains are required for fruit set, and >450 are needed for maximum seed set, with a pollen:seed ratio of about 1.6:1.0. Hummingbirds (Phaethornis superciliosus) delivered sufficient allogamous pollen for maximum seed set to only 28% of the flowers examined. Naturally pollinated flowers yielded fewer fruits and fewer seeds per fruit than those pollinated by hand. Most pollen transferred by humming-birds was self-incompatible; emasculated flowers yielded higher seed set than flowers with intact anthers. Visitation rates did not provide a good index of effective pollination.There were significant differences in ovule number, maximum seed set, and maximum per cent seed set among individual vines. More than half of an individual's flowers failed to set fruit, whether pollinated by birds or by hand. In this population, maximum reproductive potential may be limited by maternal resources for fruit development, but seed set varies with pollination intensity. Pollen-limited seed set may be a disadvantage of self-incompatibility, especially in species with many-seeded fruits.     

Floral sex allocation and reproductive success within inflorescences of Hosta ventricosa,a pseudogamous apomict

Aims Within inflorescences of sexually reproducing hermaphrodites, the production of ovules, fruits and seeds commonly declines from basal (early-opening) to distal (late-opening) flowers, while pollen production remains constant or only changes slightly, with the result that distal (late-opening) flowers become functionally male. However, few empirical studies have specifically examined whether or not changes in allocation to pollen production actually lead to changes in the number of seeds sired, a more direct measure of male fitness. In pseudogamous apomicts, fitness depends on the number of seeds produced; thus, a contrasting pattern of variation in the pollen-to-ovule (P/O) ratio within inflorescences might be expected.Methods We investigated floral sex allocation and reproductive success within racemes of Hosta ventricosa, a pseudogamous apomictic hermaphrodite possessing flowers that open acropetally. We quantified variations in pollen number, ovule number, the P/O ratio and fruit and seed production, from 2007 to 2011, among flowers within racemes of four populations of H. ventricosa in southwest China. Ecological causes for fruit and seed production were evaluated by observing patterns of pollen deposition, flower removal and supplemental pollination.Important findings Pollen number, ovule number and the P/O ratio declined from basal-to-distal positions in all sampled populations (years). Fruit and seed production decreased distally in most populations (years). Low fruit and seed set of distal flowers was not due to pollen limitation because pollen deposition never declined distally and the low fruit and seed set of distal flowers remained even after supplementary pollination was provided. The flower-removal experiment indicated that inter-fruit competition for resources among flowers was common. The low P/O ratio of distal flowers in H. ventricosa might be favored because they were unable to obtain fitness by donating pollen and siring seeds on other plants. Our study may help to understand the adaptive significance of sex allocation among flowers within inflorescences of sexually reproducing hermaphrodites.     

OVULE PACKAGING IN STOCHASTIC POLLINATION AND FERTILIZATION ENVIRONMENTS

The modular morphology of plants has important consequences for reproductive strategies. Ovules are packaged in discrete structures (flowers) that usually vary stochastically in pollen capture and ovule fertilization, because of the vagaries of pollen transfer by external agents. Different ovule packaging schemes may use limited reproductive resources more or less effectively, so that some number of ovules per flower may be optimal, given the prevailing probabilities of ovule fertilization. I derive a phenotypic model for ovule number per flower that maximizes the expected total ovule fertilizations on a plant when pollination and fertilization vary randomly among individual flowers. This model predicts that, except for small or inexpensive flowers, ovules should be “oversupplied” relative to the mean receipt of pollen tubes, so that pollen limitation of seed set should be common. Published data are congruent with this prediction. Additional hypotheses on the relation of ovule packaging to floral cost, plant size, and variance in pollen receipt are suggested by the model, but few data exist to evaluate these hypotheses.     

Active self-pollination in blackberries (Rubus subgen. Rubus, Rosaceae)

Most blackberry species are apomictic but pseudogamous, i.e. embryo development is dependent on prior pollination. Two different types of stamen arrangement were noted in senescing flowers of 18 Swedish species and 2 common cultivars. The stamens of 'cross-pollinating' species seldom touch the stigmas while those of 'self-pollinating' species instead arch over the pistils and dry up in this position. All species set seed well after artificial self-pollination and are thus self-compatible. Isolation experiments were carried out during 3 years. When inflorescences were bagged, seed set was drastically lowered in 'cross-pollinators' but remained almost unaltered in 'self-pollinators'. The different responses to isolation are probably due to the inherent stamen behaviour.     

MEGASPOROGENESIS AND GAMETOPHYTE SELECTION IN PAEONIA CALIFORNICA

Walters , James L. (U. California, Goleta.) Megasporogenesis and gametophyte selection in Paeonia californica. Amer. Jour. Bot. 49(7): 787–794. Illus. 1962.—In the ovules of Paeonia californica, a massive archesporium produces numerous (estimated at 30–40) megasporocytes, many of which complete meiosis. Several continue into gametophyte development, which is of the Polygonum type, and at the time of fertilization there are from 1 to 4 gametophytes per ovule. Rarely does more than 1 seedling per seed appear in germination. This species is characterized by extensive translocation heterozygosity, and other meiotic irregularities, in its natural populations. It shows a complete range from plants forming only pairs to those with all their chromosomes in rings at meiosis. The latter types have as high as 90% bad pollen. The course of events in the ovules is compared with the “Renner-effect” found in Oenothera. The multiple megasporocytes and subsequent events are seen as a mechanism which insures each ovule a high probability of containing a viable egg in spite of meiotic behavior which can produce 90% sterility, and thus insures high seed set in the translocation heterozygotes.     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因。为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并通过观察传粉者运动特点以及人工去花和补授花粉实验,探讨花序内资源竞争和交配环境对繁殖资源分配的影响。结果表明:(1)不同位置间的雄蕊数、雄蕊鲜重/雌蕊鲜重、花粉数及花粉胚珠比从花序基部到上部显著增加,而雌蕊鲜重和胚珠数逐渐减少,表现出上部花偏雄的性分配;上部花的结籽率显著低于基部花和中部花,不同位置间的发育种子数/果实和发育种子重/果实随着花位置的升高而显著降低,说明基部花具有更佳的雌性繁殖成效。(2)去花处理后,剩余果实的单个种子重/果实显著增加,但发育种子数/果实没有显著增加;而给上部花人工补授异花花粉后,位置间结籽率的差异消失,说明传粉限制而非资源竞争导致了花序内位置依赖的种子生产模式。(3)毛翠雀花雄性先熟的开花特征,以及传粉者苏氏熊蜂从花序基部到上部的定向访花行为,导致了花序内交配环境的变化。综上结果表明,毛翠雀花花序内的性分配和繁殖成功差异是对交配环境适应的结果,对其在高山环境中实现雌雄适合度最优化具有重要意义。     

牛耳朵传粉生物学研究

苦苣苔科(Gesneriaceae)报春苣苔属(Primulina Hance)是一个近年来备受关注的类群,其纷繁复杂的物种多样性和属下种间的特有分布引起了分类学家和植物学研究者的极大兴趣。该属除了极少数的物种如牛耳朵[(Primulina eburnea(Hance)Y. Z. Wang)]以外,绝大部分的物种为狭域分布或地方特有种,其分布范围很窄。为了揭示牛耳朵的传粉生物学和繁育系统对其生殖过程和拓殖能力的影响机制,作者系统地研究了牛耳朵的开花物候、花粉与柱头活性、访花昆虫的种类和访花行为、花粉胚珠比、OCI指数和套袋实验结实率,探究其传粉等生殖过程对牛耳朵的广布是否有正面影响。结果表明:牛耳朵的自然花期是3—5月,全花期约45 d,其中盛花期约20 d,单花期6～8 d; 开花后1～2 d花粉活力最强,开花前柱头没有可授性; 花粉胚珠比为537; 杂交指数为5; 去雌套袋、去雄套袋均无法结实,说明本种不存在无融合生殖; 与自然授粉相比,自花授粉结实率略低,异花授粉结实率略高,说明自交亲和; 牛耳朵的主要传粉者是花条蜂(Anthophora florea)和熊蜂(Bombus sp.)。花蜜产量较高、花粉量较大、花粉活力较强等特点,有利于牛耳朵完成传粉和结实的整个繁殖过程。因此,这一结果显然有利于牛耳朵的拓殖进而广布在我国华南至西南地区的喀斯特地区。     

牛耳朵传粉生物学研究

苦苣苔科(Gesneriaceae)报春苣苔属(Primulina Hance)是一个近年来备受关注的类群,其纷繁复杂的物种多样性和属下种间的特有分布引起了分类学家和植物学研究者的极大兴趣。该属除了极少数的物种如牛耳朵[(Primulina eburnea (Hance) Y. Z. Wang)]以外,绝大部分的物种为狭域分布或地方特有种,其分布范围很窄。为了揭示牛耳朵的传粉生物学和繁育系统对其生殖过程和拓殖能力的影响机制,作者系统地研究了牛耳朵的开花物候、花粉与柱头活性、访花昆虫的种类和访花行为、花粉胚珠比、OCI指数和套袋实验结实率,探究其传粉等生殖过程对牛耳朵的广布是否有正面影响。结果表明:牛耳朵的自然花期是3—5月,全花期约45 d,其中盛花期约20 d,单花期6～8 d;开花后1～2 d花粉活力最强,开花前柱头没有可授性;花粉胚珠比为537;杂交指数为5;去雌套袋、去雄套袋均无法结实,说明本种不存在无融合生殖;与自然授粉相比,自花授粉结实率略低,异花授粉结实率略高,说明自交亲和;牛耳朵的主要传粉者是花条蜂(Anthophora florea)和熊蜂(Bombus sp.)。花蜜产量较高、花粉量较大、花粉活力较强等特点,有利于牛耳朵完成传粉和结实的整个繁殖过程。因此,这一结果显然有利于牛耳朵的拓殖进而广布在我国华南至西南地区的喀斯特地区。     

Mating systems in two species ofPotentilla from Alaska

Aspects of reproductive ecology have been studied in two species ofPotentilla (Rosaceae) from interior Alaska. Data were collected to determine mating system traits such as seed: ovule and pollen: ovule ratios, and experiments were carried out in the field in order to study the effect of caging, emasculation, and manual self-pollination on seed set and seed quality. Mating system strategies differ between the two species in the populations examined. High pollen: ovule ratio, high seed: ovule ratio, good autodeposition ability, and unpredictable pollination in concert with self-incompatibility indicate thatP. hookeriana in the population studied has adopted a more or less pseudogamous mating system. In contrast, the data suggest thatP. uniflora in the studied population is approaching a sexual, facultatively outbreeding species. Because both species exhibit ploidy variation, a trait common among agamospermous plants, it is concluded that both species are likely to be facultatively agamospermous but that sexual reproduction is much more important in the population ofP. uniflora than it is inP. hookeriana.     

ARTIFICIAL INTERSPECIFIC HYBRIDIZATION IN RUELLIA (ACANTHACEAE)

Observations were made on the morphology, seed fertility, pollen viability and meiotic configurations of 22 different interspecific hybrids of 12 North American taxa representative of 3 sections of Ruellia. The hybrids were classified as (1) fully vigorous and fertile; (2) vigorous but with reduced fertility; (3) reduced vigor and fertility; (4) weak and sterile. Hybrids showed meiotic irregularities in microsporogenesis: univalents, chains, precocious separation of bivalents, nondisjunction, and micronuclei were commonly observed. Interspecific pollinations within sect. Dipteracanthus resulted in fertile or partially sterile F₁ hybrids. Intersectional crosses produced partially or completely sterile hybrids or failed. Assuming that crossability is correlated with degree of evolutionary relationship, the parental species are interpreted in genecological terms to be grouped in polytypic ecospecies. Hybridization experiments doubtless would be of value in clarifying relationships in tropical and subtropical species of Ruellia.     

Structural aspects of ovule and seed development and nonrandom abortion inMelilotus officinalis (Fabaceae)

Summary Only one ovule matures into a seed inMelilotus officinalis. Although eight ovules form within an ovary, only the basal ovule develops into a mature seed, whereas the other ovules degenerate. The investigation of ovule and seed structure at different developmental stages and a comparison of quantitative characters of differently fated ovules within an ovary were undertaken by light, phase contrast, and fluorescence microscopy. In this species, campylotropous ovules develop simultaneously on marginal placentae in an apocarpous unilocular gynoecium. Megasporo- and megagametogenesis proceed normally and are completed in bud. The maturation of the Polygonum type embryo sac takes place after the flower opens. Shortly before fertilization, synergids show signs of degeneration in all ovules. At this stage, neither the structure nor the sizes of ovules within one ovary differ significantly. In spite of this, only the basal ovule develops into a seed. Rarely, one of the upper-situated ovules or the basal and another ovule mature into seeds. Seed enlargement is insignificant until the stage when globular embryo and nuclear endosperm are formed. At the seed-filling stage, other ovules have collapsed and the seed gradually comes to occupy the total volume of the pod. The fruit-to-seed length ratio decreases considerably during seed ripening. At fertilization, ovary length is four times greater than ovule length. In the mature state, the fruit and seed lengths are approximately equal. Seed size and weight diminish with an increase in seed number within a pod, although pod size remains constant. It is assumed that nonrandom abortion of young seeds inM. officinalis is under maternal control and is not related to structural abnormalities in ovule development or with limitation in pollen. We suppose that evolution of this species may have proceeded in the direction of a decrease in seed number and an increase in its sizes, which may play an important role in seed dispersal and seedling establishment.     

Effect of pollination intensity on fruit and seed set in cacao (Theobroma cacao L.)

We studied the functional relationship between pollination intensity and fruit survival as well as the number of seeds per pod in the tropical tree Theobroma cacao L. on a Forastero Upper-Amazon clone (UPA 409) in Ivory Coast. Cutting the style 24 h after pollination allowed for counting the number of pollen grains deposited on a stigma without affecting fruit set and seed development. Forty-three pollen grains were necessary to reach 50% of maximum fruit set 28 days after pollination. Above 115 pollen grains, the proportion of developing ovaries reached a maximum of 88% 28 days after pollination and 75% at maturity. With fewer than 238 pollen grains per stigma, there was a close relationship between pollination intensity and number of seeds per pod; the pollenseed ratio increased from 1.61 to 3.81 for PI increasing from 30 to 238 pollen grains. For higher pollination intensities, the average number of seeds per pod reached a maximum of 58. The relationship between pollination intensity and seed content was modelled. Results are consistent with the hypothesis that ovules attracted pollen tubes in a similar way regardless of whether or not they had already been reached by another pollen tube.     

菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系

三叶鬼针草(Bidens pilosa)是一种危害严重的菊科入侵植物。通过实验观察和人工控制套袋等方法, 对其花序开花动态、花粉胚珠比 (P/O)、自交亲和性、花粉活力、访花昆虫和种子 (瘦果)的萌发率等与繁殖相关的特征进行了研究, 探讨了这些繁殖特征与入侵性的关系。三叶鬼针草在10-11月开花, 单个花序的花期约为5-6天。每小花的花柱基部有圆筒状的蜜腺环绕。单个花序内可自交亲和, 自交结实率和花粉活力均较高, 其P/O值为1 7 54.1 2±29 .8 7。主要访花者为灰蝶科(Lycaenidae)、粉蝶科(Pieridae)和茧蜂科(Braconidae)昆虫。三叶鬼针草所具有的灵活交配机制是其入侵成功的重要因素。此外, 三叶鬼针草结实量大、种子产生迅速且适于传播, 以及种子萌发范围广和短期快速萌发等特性也增强了其入侵性。