冠果草的性表达状态及其进化含义

通常认为雄花两性花同株是植物性表达两性花进化到雌雄异株的过渡类型之一。慈姑属仅冠果草（Ｓａｇｉｔｔａｒｉａ ｇｕｙａｎｅｎｓｉｓ Ｈ．Ｂ．Ｋ．ｓｕｂｓｐ．ｌａｐｐｕｌａ（Ｄ．Ｄｏｎ）Ｂｏｊｉｎ）的性表达为典型的雄花两性花同株,其他种为雌雄同株。对冠果草野外和实验居群的性配置进行了定量观察。东乡和武夷山居群雄花的比例分别为２．４８％０．９６％。实验表明,栽培条件影响冠果草的性表达：在营养较好的条件下     

Subandrodioecy and male fitness in Sagittaria lancifolia subsp. lancifolia (Alismataceae)

Sagittaria lancifolia subsp. lancifolia is described as cosexual (monoecious), but the study population consisted of 84% cosexuals that typically had 35% pistillate buds and 16% predominant males that typically had 0-2% pistillate buds. Hand-pollinations showed that pistillate flowers required pollination to set seed, and pollen from both male and cosexual plants was potent. No gender switching was seen in the field or greenhouse. From 24 experimental crosses, 890 offspring were grown to maturity. Among these, all offspring of cosexual sires were cosexual, but approximately half the offspring of male sires were male, implying that maleness was inherited as a single, dominant allele. These results indicate that S. lancifolia is subandrodioecious, a very rare breeding system. It is rare, in part because its maintenance requires a large male-fitness differential between male and cosexual plants. In the study population, this condition was met by the differential survival of staminate buds on male racemes. Larvae of the weevil Listronotus appendiculatus killed many staminate buds. They did so in a vertical gradient, with buds lower on racemes safer. Male plants have replaced pistillate with staminate buds at these safer positions and thereby enjoy disproportionally higher male fitness.     

An Examination of the Function of Male Flowers in an Andromonoecious Shrub Capparis spinosa

The pollen donor and pollinator attractor hypotheses are explanations for the functions of the male flowers of andromonoecious plants. We tested these two hypotheses in the andromonoecious shrub Capparis spinosa L. （Capparaceae） and confirmed that pollen production and cumulative volume and sugar concentration of nectar do not differ between male and perfect flowers. However, male flowers produced larger anthers, larger pollen grains and smaller ovaries than perfect flowers. Observations on pollinators indicated that two major pollinators （Xylocopa valga Gerst and Proxylocopa sinensis Wu） did not discriminate between flower morphs and that they transferred pollen grains a similar distance. However, there were more seeds per fruit following hand pollination with pollen from male flowers than from perfect flowers. Individuals of C. spinosa with a larger floral display （i.e. bearing more flowers） received more pollen grains on the stigma of perfect flowers. Female reproductive success probably is not limited by pollen. These results indicate that male flowers of C. spinosa save resources for female function and that they primarily serve to attract pollinators as pollen donors.     

Embryological Studies on Sagittaria guayanensis H. B. K. Subsp. lappula (D. Don) Bojin (Alismataceae)

This paper deals with the embryological characteristics of Sagittaria guayanensis H. B.K. subsp. lappula (D. Don) Bojin. The anther wall development follows the Monocotyledonous type. The cytokinesis of microspore mother cell in meiosis is of the Successive type. The tetrads of microspores show an isobilateral arrangement, and the mature pollen grains are 3-celled. The ovule is bitegminous, pseudo-crassinucellate and anatropous. The megaspore mother cell originates directly from a single archesporial cell. The mature embryo sac consists of 7 cells including 8 nuclei and conforms to the Allium type. The two polar nuclei do not fuse into a secondary nucleus before fertilization. Instead, one sperm fuses with the micropylar end polar nucleus first , and the fertilized polar nucleus then migrates to the chalazal end, where it fuses with the second polar nucleus, forming the primary endosperm nucleus. The embryo development conforms to the Caryophyllad type. The mature embryo is U-shaped and forms the embryonic shoot apex accompanied by two leaves. The endosperm development corresponds to the Helobial type. The primary endosperm nucleus (invariably lying in the chalazal part of the embryo sac) divides and forms two chambers:large micropylar one and small chalazal one. The chalazal endosperm chamber remains binucleate, while, in the micropylar chamber free nuclear divisions occur and then cellnlarization takes place. During the embryo formation the endosperm gradually degrades and can not be found in the mature seed. The subgenus Lophotocarpus is different from the subgenus Sagittaria in some embryological aspects, especially in the structure of mature embryo sac and the double fertilization process.     

An Examination of the Function of Male Flowers in an Andromonoecious Shrub Capparis spinosa

The pollen donor and pollinator attractor hypotheses are explanations for the functions of the male flowers of andromonoecious plants. We tested these two hypotheses in the andromonoecious shrub Capparis spinosa L. (Capparaceae) and confirmed that pollen production and cumulative volume and sugar concentration of nectar do not differ between male and perfect flowers. However, male flowers produced larger anthers, larger pollen grains and smaller ovaries than perfect flowers. Observations on pollinators indicated that two major pollinators (Xylocopa valga Gerst and Proxylocopa sinensis Wu) did not discriminate between flower morphs and that they transferred pollen grains a similar distance. However, there were more seeds per fruit following hand pollination with pollen from male flowers than from perfect flowers. Individuals of C. spinosa with a larger floral display (i.e. bearing more flowers) received more pollen grains on the stigma of perfect flowers. Female reproductive success probably is not limited by pollen. These results indicate that male flowers of C. spinosa save resources for female function and that they primarily serve to attract pollinators as pollen donors.     

The evolution and maintenance of monoecy and dioecy in Sagittaria latifolia (Alismataceae)

Plant species rarely exhibit both monoecious and dioecious sexual systems. This limits opportunities to investigate the consequences of combined versus separate sex function on mating patterns and genetic variation and the analysis of factors responsible for the evolution and maintenance of the two sexual systems. Populations of the North American clonal aquatic Sagittaria latifolia are usually either monoecious or dioecious and often grow in close geographic proximity. We investigated mating patterns, genetic structure, and relationships between the two sexual systems using allozyme variation in populations from southern Ontario, Canada. As predicted, selfing rates in monoecious populations (n = 6, mean = 0.41) were significantly higher than in dioecious populations (n = 6, mean = 0.11). Moreover, marker-based estimates of inbreeding depression (delta) indicated strong selection against inbred offspring in both monoecious (mean delta = 0.83) and dioecious (mean delta = 0.84) populations. However, the difference in selfing rate between the sexual systems was not reflected in contrasting levels of genetic variation. Our surveys of 12 loci in 15 monoecious and 11 dioecious populations revealed no significant differences in the proportion of polymorphic loci (P), number of alleles per locus (A), or observed and expected heterozygosity (H(o) and H(e), respectively). Strong inbreeding depression favoring survival of outcrossed offspring may act to maintain similar levels of diversity between monoecious and dioecious populations. Despite geographical overlap between the two sexual systems in southern Ontario, a dendrogram of genetic relationships indicated two distinct clusters of populations largely corresponding to monoecious and dioecious populations. Reproductive isolation between monoecious and dioecious populations appears to be governed, in part, by observed differences in habitat and life history. We suggest that selfing and inbreeding depression in monoecious populations are important in the transition from monoecy to dioecy and that the maintenance of distinct sexual systems in S. latifolia is governed by interactions between ecology, life history, and mating.     

Population genetics of two rare perennials in isolated wetlands: Sagittaria isoetiformis and S. teres (Alismataceae)

We investigated genetic structure in two closely related perennial plants that occur in isolated wetlands: Sagittaria isoetiformis, restricted to the southeastern Coastal Plain of North America, and S. teres, endemic to the northeastern Coastal Plain. Using horizontal starch-gel electrophoresis, we screened 527 individuals from 11 populations of S. isoetiformis and 367 individuals from seven populations of S. teres. A high proportion of the 16 loci were polymorphic (%P(S) = 93.8% in S. isoetiformis and %P(S) = 75.0% in S. teres), with higher mean numbers of alleles per polymorphic locus and effective alleles per locus in S. isoetiformis (AP = 3.27, A(E) = 1.90) than in S. teres (AP = 2.58, A(E) = 1.30). Species- and population-level expected heterozygosities were higher in S. isoetiformis (H(ES) = 0.399, H(EP) = 0.218) than in S. teres (H(ES) = 0.177, H(EP) = 0.101). Jackknife estimates of F statistics indicated moderate levels of inbreeding in S. teres (F(IS) = 23.1%). Strong differentiation characterized these geographically isolated populations (G(ST) = 39.9% in S. isoetiformis, and G(ST) = 26.1% in S. teres). Genetic identities varied substantially within (ī = 75%, range = 0.558-0.963 in S. isoetiformis; ī = 89%, range = 0.776-0.963 in S. teres) and among species (ī = 81%, range = 0.506-0.882), leading to the discrimination of four regional population clusters using nonmetric multidimensional scaling (NMDS). It appears that S. isoetiformis and S. teres are a progenitor-derivative species pair.     

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae)

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.     

Fruit-Set Induced Changes in the Sex of Flowers in Caesalpinia calycina (Leguminosae)

Abstract: The proportions of hermaphrodite to staminate flowers in inflorescences of the andromonoecious species Caesalpinb calycina vary. Analysis of fruit position along the flowering rhachis, and also of the sex of floral buds in inflorescences with fruits set in different positions, indicate that fruiting success influences the sex of flowers in distal positions along the rhachis. Other reports of andromonoecy in caesalpinoid species are examined with reference to floral sex lability and the influence of fruit set.     

Sex determination and the evolution of dioecy from monoecy in Sagittaria latifolia (Alismataceae)

The role of mutations of small versus large effect in adaptive evolution is of considerable interest to evolutionary biologists. The major evolutionary pathways for the origin of dioecy in plants (the gynodioecy and monoecy-paradioecy pathways) are often distinguished by the number of mutations involved and the magnitude of their effects. Here, we investigate the genetic and environmental determinants of sex in Sagittaria latifolia, a species with both monoecious and dioecious populations, and evaluate evidence for the evolution of dioecy via gynodioecy or monoecy-paradioecy. We crossed plants of the two sexual systems to generate F1, F2 and backcross progeny, and grew clones from dioecious populations in low-and high-fertilizer conditions to examine sex inconstancy in females and males. Several lines of evidence implicate two-locus control of the sex phenotypes. In dioecious populations sex is determined by Mendelian segregation of alleles, with males heterozygous at both the male- and female-sterility loci. In monoecious populations, plants are homozygous for alleles dominant to male sterility in females and recessive to female sterility in males. Experimental manipulation of resources revealed sex inconstancy in males but not females. These results are consistent with predictions for the evolution of dioecy via gynodioecy, rather than the expected monoecy-paradioecy pathway, given the ancestral monoecious condition.     

利用腊叶标本初探豹子花属植物的性表达及其与个体大小的关系?

植物性系统进化过程中很少有植物通过雌性败育的方式实现性系统由联合走向分离,然而雄花在百合科植物中大量存在,引起了研究者的关注。本研究利用百合科豹子花属植物的腊叶标本,获取它们的花性别表型、花大小、株高以及分布海拔等信息探讨豹子花属植物的性表达样式,并对样本量较多的开瓣豹子花、云南豹子花、多斑豹子花和豹子花四个种的性表达及其与个体大小的关系进行分析。研究结果表明：1)六种豹子花属植物标本中都有雄花存在;根据雄性花和两性花的不同组合,形成雄性个体、两性花个体,和雄花两性花同株个体三种植株类型;2)个体大小(株高)和分布海拔之间不存在相关关系,和植株性别类型相关,即雄性个体一般显著小于具有雌性功能的两性花个体和雄花两性花同株个体;3)个体大小与花大小、每个个体开花总数和雌花数量成显著的正相关关系,而与每个个体的雄花数量无显著的相关关系。本研究更正了一直以来对豹子花属植物性系统的错误认识,首次报道雄花在豹子花属植物中普遍存在,并指出雄花和雄性个体的出现是大小依赖性分配的结果。同时,以研究实例证明了利用腊叶标本开展植物性系统多样性的可行性。     

Effects of different nutrient sources on plasticity of reproductive strategies in a monoecious species,Sagittaria graminea (Alismataceae)

Varied nutrient sources can influence the plasticity of reproductive strategies in monoecious species differently. We examined the plasticity of sexual and clonal reproductive components in distinct nutrient sources in a monoecious species, Sagittaria graminea Michx. The results showed that for aboveground reproductive components, in rich-phosphorus and low-nitrogen conditions, the species produced more male flowers, whereas female flowers did not significantly increase in number compared to low-phosphorus and low-nitrogen conditions (control). In rich-nitrogen and low-phosphorus conditions, the species produced more flowers, particularly female, and more seeds, compared to the control conditions. In the rich-nitrogen and rich-phosphorus conditions, plants increased male flowers, female flowers, total flower number, synchronously, as well as seed production. For the belowground reproductive components, in the rich-phosphorus conditions, the plants produced bigger corms than in the rich-nitrogen and control conditions, which further enhanced their competitiveness against companion species. In rich-nitrogen conditions, the plants produced more medium and small corms, and relatively more and longer stolons, which were useful in expansion and invasion of more space. Furthermore, the species displayed trade-off relationships between the aboveground and belowground reproductive components in unbalanced nutrient conditions (add-N or add-P). However, in low nutrient levels (control) or in nutrient abundant (add-N + P) conditions, there were no significant trade-off relationships between the aboveground and belowground reproductive components, based on quantities.     

Pattern of Flower and Fruit Production in Stryphnodendron adstringens, an Andromonoecious Legume Tree of Central Brazil

Abstract: Patterns of flower and fruit production in racemes of Stryphnodendron adstringens, an andromonoecious Brazilian savanna tree species, were studied in two natural areas near Uberlândia-MG. Racemes were divided in three parts: apex, centre, and base. Number of flowers, gender, and nectar and pollen production were analyzed for each section. Frequency of visitors to each part of the inflorescence was also quantified. Hand self- and cross-pollinations were performed in complete racemes and fruit set used to determine breeding system. The racemes produced a mean of 329 flowers, more densely packed in the central portion. Hermaphrodite and male flowers occur along the inflorescence but hermaphrodite flowers are more common in the centre. Fruit set was markedly low but does not seem to be limited by pollination service, since free open-pollinated racemes and hand cross-pollinated ones do not differ in fruit production rates. Fruits resulted mostly from cross-pollinated flowers and fruit production was biased to the central portion of the raceme. Nectar yield was higher in the central portion of the raceme and visitors arrived more commonly on this portion of the inflorescence. However, most flowers did not produce nectar. The pattern of fruit production seems to be a consequence of the hermaphrodite flower distribution in the raceme and it is not constrained by pollen flow or flower opening sequence.     

Phenotypic plasticity of hermaphrodite sex allocation promotes the evolution of separate sexes: an experimental test of the sex-differential plasticity hypothesis using Sagittaria latifolia (Alismataceae)

Separate sexes can evolve under nuclear inheritance when unisexuals have more than twice the reproductive fitness of hermaphrodites through one sex function (e.g., when females have more than twice the seed fertility of hermaphrodites). Because separate sexes are thought to evolve most commonly via a gynodioecious intermediate (i.e., populations in which females and hermaphrodites cooccur), the conditions under which females can become established in populations of hermaphrodites are of considerable interest. It has been proposed that resource-poor conditions could promote the establishment of females if hermaphrodites are plastic in their sex allocation and allocate fewer resources to seed production under these conditions. If this occurs, the seed fertility of females could exceed the doubling required for the evolution of unisexuality under low-, but not high-resource conditions (the sex-differential plasticity hypothesis). We tested this hypothesis using replicate experimental arrays of the aquatic herb Sagittaria latifolia grown under two fertilizer treatments. The results supported the sex-differential plasticity hypothesis, with females having more than twice the seed fertility of hermaphrodites under low-, but not high-fertilizer conditions. Our findings are consistent with the idea that separate sexes are more likely to evolve under unfavorable conditions.     

The effects of density on size-dependent gender plasticity in the monoecious species Sagittaria potamogetifolia (Alismataceae)

Aim

To test the fitness-gain curve model proposes that cosexual plants adjust their sex ratios and resource allocation depending on their size. In this study, the monoecious species Sagittaria potamogetifolia was used as a model to determine the effects of plant size and density on gender modification and reproductive allocation.

Bacterial (Bacillus thuringiensis subsp. thuringiensis and B. thuringiensis subsp. kurstaki) Entomocidal Preparations Decrease Chlorophyll Accumulation in Larch Needles

Chlorophyll a plus b content and absorption spectra of the homogenates from the cotyledonary leaves of 30-day-old seedlings of two larch species, Larix gmelinii (Rupr.) Rupr. and L. sibirica Ldb. were studied. The seedlings were grown on Perlite containing aqueous solutions of entomocidal biopreparations isolated from Bacillus thuringiensis subsp. thuringiensis (bitoxybacillin) and B. thuringiensis subsp. kurstaki (lepidocide) at various final concentrations (2, 6, and 12 g/l). Changes in the form of chlorophyll absorption spectra induced by biopreparations were established. A marked inhibition of pigment accumulation in the needles dependent on the biopreparation concentration was noted. At a low concentration (2 g/l), the biopreparations virtually did not affect the chlorophyll content; an increase in their concentrations resulted in a decrease in chlorophyll content in leaves by 20% (at 6 g/l) and 40% (at 12 g/l). It is concluded that bitoxybacillin and lepidocide inhibited the chlorophyll accumulation in larch needles to a similar extent.     

Reproductive biology of the andromonoecious Cucumis melo subsp. agrestis (Cucurbitaceae)

Background and Aims

Cucumis melo subsp. agrestis (Cucurbitaceae) is cultivated in many African regions for its edible kernels used as a soup thickener. The plant, an annual, andromonoecious, trailing-vine species, is of high social, cultural and economic value for local communities. In order to improve the yield of this crop, the first step and our aim were to elucidate its breeding system.

Methods

Eight experimental pollination treatments were performed during three growing seasons to assess spontaneous selfing, self-compatibility and effects of pollen source (hermaphroditic vs. male flowers). Pollination success was determined by pollen tube growth and reproductive success was assessed by fruit, seed and seedling numbers and characteristics. The pollinator guild was surveyed and the pollination distance determined both by direct observations and by indirect fluorescent dye dispersal.

Key Results

The species is probably pollinated by several Hymenoptera, principally by Hypotrigona para. Pollinator flight distances varied from 25 to 69 cm. No evidence for apomixis or spontaneous self-pollination in the absence of insect visitors was found. The self-fertility index (SFI = 0) indicated a total dependence on pollinators for reproductive success. The effects of hand pollination on fruit set, seed number and seedling fitness differed among years. Pollen tube growth and reproductive success did not differ between self- and cross-pollinations. Accordingly, a high self-compatibility index for the fruit set (SCI = 1·00) and the seed number (SCI = 0·98) and a low inbreeding depression at all developmental stages (cumulative δ = 0·126) suggest a high selfing ability. Finally, pollen origin had no effect on fruit and seed sets.

Conclusions

This andromonoecious species has the potential for a mixed mating system with high dependence on insect-mediated pollination. The selfing rate through geitonogamy should be important.     

Sex ratio and reproductive effort in the dioecious Juniperus communis subsp. alpina (Suter) Celak. (Cupressaceae) along an altitudinal gradient

The hypothesis that reproductive cost differs between sexes was tested in Juniperus communis subsp. alpina along an altitudinal gradient. Sex ratio (male : female) increased significantly with elevation, and above 2,600 m it was significantly male-biased. The reproductive effort was markedly greater for females than for males at all elevations. However, over 3 years of study, the growth of the females, measured as elongation of the main axes, was similar to that of the males. In both sexes, growth decreased with increasing elevation. Neither size of the ripe seed cones, nor the number of developed seeds per cone varied with elevation. The percentage of filled seeds was significantly greater at higher elevations indicating more favourable conditions for wind pollination in these stands. However, cone production decreased with elevation and so, reproductive success of J. communis subsp. alpina in Sierra Nevada decreases towards both upper and lower altitudinal distribution limits. The results do not support the hypothesis of differential reproductive cost between sexes; thus, alternative arguments to explain the altitudinal variation of sex ratio are discussed.     

H.ylori尿素酶B亚单位在保加利亚乳杆菌的表达与鉴定

目的构建表达幽门螺杆菌（Helicobacter pylori,H、pylori）尿素酶B亚单位（UreB）的基因工程乳杆菌,并对其进行初步的安全性评估。方法采用高保真PCR从H.pylori标准菌株NCTC 11637中扩增ureB基因,插入乳酸菌表达质粒pMG36e,将重组质粒电转入保加利亚乳杆菌L6032中,获得表达ureB的基因工程乳杆菌。在含乳糖的MRS培养基诱导目的蛋白表达,Western blot鉴定其免疫原性。连续传代培养60代,检测基因工程乳杆菌的稳定性、形态学与生理生化特性以进行初步的安全性评估。结果特异PCR、酶切和测序鉴定均证实ureB基因克隆入表达载体pMG36e,SDS-PAGE结果显示,重组质粒pMG36e-ureB电转入保加利亚乳杆菌所构建的基因工程乳杆菌能表达约64KD的蛋白,Western blot证明该蛋白能与抗H.priori ureB的兔血清反应。稳定性、形态学与生理生化特性检测结果表明,基因工程乳杆菌与原始菌株保加利亚乳杆菌完全一致。结论成功构建能表达H.pylori UreB的保加利亚乳杆菌L6032-UreB,该基因工程菌在形态与生理生化特性上未发生任何变异,从而为探索幽门螺杆菌感染的益生菌制剂调理疗法奠定了坚实的基础。