Analysis of late stage flower development in Primula vulgaris reveals novel differences in cell morphology and temporal aspects of floral heteromorphy

Heterostyly in Primula is characterized by the development of long-styled pin and short-styled thrum flowers, with anthers midway down the corolla tube in pin flowers, and at its mouth in thrum flowers. Other differences include pollen size and stigmatic papillae length. Several linked genes at the S locus control these differences.In this study we have analyzed pin and thrum flowers through the temporal development of heteromorphy.These studies indicate that the S locus linked genes that orchestrate heteromorphic flower development act in coordination, but with different temporal and spatial dynamics. Style length is differentiated by longer style cells in pin than thrum. However, our studies on cell shape and size within the corolla tube show that a different mechanism mediates the dissimilar elevation of anthers between pin and thrum types. These studies have also revealed that upper corolla tube cells in thrum flowers are wider than those in pin flowers. This results in a larger corolla tube mouth in thrum flowers and represents a new and previously undocumented heteromorphic variation between pin and thrum flowers.     

Are any primroses (Primula) primitively monomorphic?

Primula (c. 430 species) and relatives (Primulaceae) are paradigmatic to our understanding of distyly. However, the common co-occurrence of distyly and monomorphy in closely related groups within the family has made the interpretation of its evolution difficult.Here, we infer a chloroplast DNA (cpDNA) phylogeny for 207 accessions, including 51% of the species and 95% of the sections of Primula with monomorphic populations, using Bayesian methods. With this tree, we infer the distribution of ancestral states on critical nodes using parsimony and likelihood methods.The inferred cpDNA phylogeny is consistent with prior estimates. The most recent common ancestor (MRCA) of Primula is resolved as distylous using both methods of inference. However, whether the distyly in Primula, Hottonia, and Vitaliana arose once or three independent times is not clear.We conclude that monomorphism in descendants of the MRCA of Primula is derived from distyly in all cases. Thus, scenarios for the evolution of distyly that rely on the persistence of primitive monomorphy (such as in Primula section Sphondylia) require re-evaluation.     

Pollen flow in a population of Primula vulgaris Huds.

A British population of Primula vulgaris was visited four times between 1971 and 1976 for study of the nature and functioning of distyly. The stainability and production of pollen from pins exceeded those of thrums. About 20% of pin pollen and 48% of thrum pollen was removed from anthers by insect visitors. Pins and thrums were present in a 1:1 ratio in 1971 but pins outnumbered thrums in 1976. Thrums produced more seeds per flower than pins though the number of ovules produced by each was similar. Two methods for collecting stigmas from open flowers and analyzing them for pollen loans produced somewhat different results. For pin stigmas, the pollen load consisted of 2–23% thrum pollen; for thrum stigmas, the pollen load consisted of 0–71% pin pollen with most stigmas having less than 50% pin pollen. In general, intermorph pollen flow is less than would be expected if pollen flow were random. It is probable that most intramorph pollen on stigmas is a result of self- or geitonogamous pollination. The extensive literature concerning the natural pollinators of the primrose is reviewed. Although Darwin's hypothesis concerning the functional significance of distyly in promoting intermorph pollination was never quantified, the pollen flow patterns observed in P. vulgaris are unexpectedly deviant and are similar to those patterns observed in several unrelated heterostylous species in other plant families.     

Pollen Morphology and Taxonomic Position of the Genus Pentaphragma Wall. (Pentaphragmataceae)

The pollen morphology of some Pentaphragma Wall, species has been studied by light- and electron microscopy. To improve the rehydration of the dried material used, compound fixatives have been applied. Intine-held vesicles and fibrillar network in gaps and spaces of the exine stain positively for protein. The pollen morphology of the genus differs strongly from that of Campanulaceae, and shows distinctive characters of its own. Hence from a palynological point of view it appears correct to place the genus in a separate family.     

Evolution of syncarpy in angiosperms: theoretical and phylogenetic analyses of the effects of carpel fusion on offspring quantity and quality

The repeated evolution of fused carpels (syncarpy) is one of the dominant features of angiosperm macroevolution. We present results of new phylogenetic and theoretical analyses to assess the frequency and nature of transitions to syncarpy, and the possible advantages of syncarpy over apocarpy under a variety of ecological conditions. Using a recent molecular estimate of angiosperm phylogeny, we ascertained that a minimum of 17 independent evolutionary transitions from apocarpy to syncarpy have occurred; about three‐quarters of these transitions allowed pollen tubes to cross between carpels and fertilize ovules that would otherwise be left unfertilized. Most of these transitions also intensified competition between pollen, potentially enhancing offspring fitness. The high proportion of evolutionary transitions promoting pollen competition and pollen‐tube access to all carpels supports the hypothesis that the main advantage of syncarpy is in increasing offspring quality and quantity. The potential advantages of syncarpy were more thoroughly evaluated by analytical and simulation studies. These showed that the advantage of syncarpy over apocarpy involving increased offspring‐quantity held under conditions of marginal pollination and declined with increasing pollination. The offspring‐quality advantage persisted over a wider range of conditions, including under quite high pollination rates.     

The distyly in Primula elatior (L.) Hill (Primulaceae), with a study of flowering phenology and pollen flow

Flowers of Primula elatior exhibit all the dimorphic traits typical of distyly. The difference in pollen production between pin and thrum flowers is not thought to be caused by the pollen size dimorphism. The percentage of pin pollen which is probably viable exceeds that of thrum pollen. In the population studied the morph ratio is 1:1, and plants are randomly dispersed with regard to morph type. Bumblebees are the main pollinators, and analysis of the stigmatic pollen load in naturally |xjllinated flowers shows that pin stigmas receive many more pollen grains than thrum stigmas. Intramorph pollination is considerable in pins, while thrums are subject to some differential pollination. Emasculation experiments have shown that intraflower pollination in the thrum morph is negligible. Aging of pin flowers has a reducing effect on their total pollen loads, while in thrums there is no change in total pollen load after the first days of anthesis.     

胭脂花野生居群花粉形态多样性研究

胭脂花是分布于中国北部的报春花科报春花属花柱二型植物,为了阐明胭脂花花粉形态的多样性,对胭脂花分布范围内的7个野生居群的花粉形态进行了测定。结果表明,胭脂花花粉属N3P4C5型,极面观多为三角形,具三孔沟,少数为四边形,具4孔沟;短花柱植株的花粉粒较大(16.53×17.58μm),花粉粒外壁粗糙,表面纹饰多,网眼大;长花柱植株的花粉粒较小(12.95×13.72μm),花粉粒外壁较光滑,表面纹饰少,网眼小。不同居群间花粉粒在大小、网眼直径、外壁纹饰等方面存在显著差异,花粉形态变异系数在4.88%~35.48%之间;短花柱类型各居群的平均变异系数为16.18%,居群间差异较大,其中庞泉沟居群变异系数最大(24.05%),海坨山居群最小(10.13%);长花柱类型各居群的平均变异系数为14.89%,庞泉沟居群的变异系数最大(19.75%),五台山与海坨山居群最小(12.50%),长花柱类型的花粉性状比短花柱植株更为稳定。UPGMA聚类分析发现,7个居群聚为两大类,除庞泉沟居群外,其他居群聚为一组,地理距离相近的居群一般聚在一起。研究结果为胭脂花种质资源遗传多样性及育种研究提供了依据。     

Phylogenetic relationships and character evolution in Primula L.: The usefulness of ITS sequence data

ABSTRACT

The main goals of this research were to reconstruct the infrageneric phylogeny of the genus Primula based on both nuclear and chloroplast DNA sequences, and to use the resulting phylogenies to elucidate the evolution of breeding systems, morphological characters, chromosome number, and biogeographic distribution in the genus. In this paper, the results of a pilot study based on the nuclear ribosomal Internal Transcribed Spacer (ITS) region are described. ITS sequences from 21 taxa produced a number of variable characters sufficient to resolve relationships among sections. The resulting phylogeny confirmed the monophyly of sections Auricula and Aleuritia. Sections Armerina, Proliferae, Crystallophlomis, Parryi, and Auricula, with a base chromosome number of x = 11, and sect. Aleuritia, with a base chromosome number of x = 9, formed two well supported clades. The ITS topology also suggested that leaves with revolute vernation, previously believed to be a derived state, might represent the ancestral condition in Primula, with later reversals to the involute condition. Finally, this initial ITS tree provides preliminary support to the proposed role of the widespread, diploid and heterostylous P. mistassinica as having given origin to the polyploid, homostylous P. incana and P. laurentiana.     

Palynological study of Veronica Sect. Veronica and Sect. Veronicastrum (Scrophulariaceae) and its taxonomic significance

Pollen grains from 30 taxa, mainly from the Western Mediterranean area belonging to the perennial sections of the genus Veronica (Scrophulariaceae), were studied by LM and SEM. A close relationship was found between pollen size and ploidy level. Palynological characters cannot be regarded as essential taxonomic tools within the groups studied, although they have systematic value in some specific cases. The monophyly of Veronica subsect. Multiflorae is supported by a common and exclusive sculpture of exine. Delimitation of the genus is discussed and hypothetical evolutionary trends concerning pollen characters are addressed.     

中国特有濒危种川东灯台报春(报春花科)的重新发现

2006年6月份和9月份,作者在对重庆市城口县大巴山地区进行考察时,采集到了中国特有种川东灯台报春（Primula mallophylla）的标本,这是近百年来对该种除模式标本外的再度发现。根据在大巴山观察及采集到的材料,作者对过去该种的特征描述中存在的一些疑问进行了澄清,更正和补充了其形态描述;对历史上混淆的几个种列出了检索表;报道了该种的染色体数目及核型;根据国际通用的“红色名录等级及标准”对其濒危等级进行了评估。     

木里报春一新异名

将新近发表的Primula hongshanensis D. W. H. Rankin, Z. D. Fang &; H. Sun处理为木里报春P. boreiocalliantha的异名, 对近缘的几个种列出了检索表。