蕨类植物性别分化对环境的响应

蕨类植物是维管植物中唯一的孢子体和配子体都能独立生活的类群.同型孢子蕨类配子体的性别分化受到激素和环境因子的影响.生理学研究表明,成精子囊素与赤霉素能诱导雄配子体发育,抑制雌配子体发育;脱落酸阻止成精子囊素诱导的精子器形成;乙烯合成前体ACC促进赤霉素诱导的精子器形成,而乙烯合成抑制因子AOA通过抑制细胞分化来抑制精子器形成.光照对不同种类蕨类配子体分化的影响存在差异.糖类能够促进雄配子体形成,并可加速成熟雌配子体向两性分化.钙离子、钴离子和甲硫氨酸等分别参与了蓝光和赤霉素对配子体性别分化的调控过程.培养密度影响配子体生长及性别表达,高密度下雄性和无性配子体居多,而低密度下两性和雌性配子体居多.近年来的突变体表型分析与分子生物学研究表明,成精子囊素通过影响ANI1、HER、TRA、FEM和MAN等基因的表达调控配子体性别分化.综述了蕨类植物性别分化对环境响应的研究进展.     

麦秆蹄盖蕨（Athyrium fallaciosum）配子体的发育规律及播种密度对配子体性别分化的影响

在组织培养条件下,对麦秆蹄盖蕨（Athyrium fallaciosum）配子体发育的连续过程进行了详细观察。结果表明:麦秆蹄盖蕨孢子为四面体型; 孢子萌发为书带蕨型（Vittaria-type）;原叶体发育为铁线蕨型（Adiantum-type）,成熟原叶体为对称的心形;精子器近圆球形,成熟颈卵器细长,常向原叶体基部倾斜或弯曲。常规播种条件下,发现麦秆蹄盖蕨配子体有雌配子体、雄配子体、雌雄同体配子体和无性配子体类型。配子体的性别随密度不同而呈现一定的变化趋势,雄配子体随密度增大呈上升趋势;雌配子体随密度增大先上升后下降;雌雄同体配子体和无性配子体随密度变化不大。雌配子体和雌雄同体配子体具颈卵器数目一般为10～15个;精子器数目随密度的增大逐渐减少,雄配子体中具有约50个精子器,雌雄同体配子体具有约20个精子器。     

Growth and gender in the gametophyte of Blechnum spicant L

The growth and gender in the gametophyte of Blechnum spicant L. were strongly affected by its origin, spore or mature homogenized gametophytes, and also by the addition of plant growth regulators to the culture medium. Spore-derived gametophytes cultured in Murashige and Skoog liquid medium were females and heart-shaped, whereas those derived from mature homogenized gametophytes were shorter and male or asexual (1:3) because of the release of antheridiogen to the culture medium. In the latter, maleness was especially increased by the addition of BA. This cytokinin is a strong inductor of maleness in homogenized cultures; however, even though BA influenced sexual organ formation in spore-derived gametophytes, it does not change the␣female sexual pattern that these gametophytes have. Separate male and female populations of gametophytes were obtained in this work.     

Growth and reproductive characteristics in artificially formed clonal gametophytes ofDryopteris filix-mas (Dryopteridaceae)

Artificially formed clonal gametophytes ofDryopteris filix-mas (L.)Schott were able to grow and reproduce, although growth rates, proportions of hermaphrodites and reproductive efforts were low. Variable density and nutrient levels appeared to affect gametophytic life histories and they continued to influence the viability of the developing sporophyte generation. Differences between populations and sporophytes were discovered in life histories. Hermaphroditic gametophytes were considerably larger and possessed higher viability than did males. Clear differences between clones in the level of phenotypic plasticity in gametophyte size were observed. No neighbor effect on gametophyte sex expression was detected with the densities and nutrient levels used.     

Comparison of gametophyte growth, sex determination and reproduction in three fern species from the tropics

The gametophytes of Athyrium boryanurn, Thelypteris subpubescens and Pteridium aquilinum (vars. aquilinum, esculentum and wightianum) were grown as isolates, pairs and groups, and they were found to differ in their life histories: A. boryanum is a selfer which is sensitive to competition, while T. subpubescens is an ineffective selfer which is sensitive to competition, and P. aquilinurn is generally ineffective at sexual reproduction. A relationship between the level of gametophyte reproduction and the sporophyte viability was detected. The interactions between neighbours (competition or antheridiogen effects) appeared to influence gametophyte growth and reproductive success. In general, hermaphroditic gametophytes were considerably larger than males in all species, and those hermaphrodites which formed sporophytes were often smaller than other hermaphrodites.     

Sexual conflict and the alternation of haploid and diploid generations

Land plants possess a multicellular diploid stage (sporophyte) that begins development while attached to a multicellular haploid progenitor (gametophyte). Although the closest algal relatives of land plants lack a multicellular sporophyte, they do produce a zygote that grows while attached to the maternal gametophyte. The diploid offspring shares one haploid set of genes with the haploid mother that supplies it with resources and a paternal haploid complement that is not shared with the mother. Sexual conflict can arise within the diploid offspring because the offspring's maternal genome will be transmitted in its entirety to all other sexual and asexual offspring that the mother may produce, but the offspring's paternally derived genes may be absent from these other offspring. Thus, the selective forces favouring the evolution of genomic imprinting may have been present from the origin of modern land plants. In bryophytes, where gametophytes are long-lived and capable of multiple bouts of asexual and sexual reproduction, we predict strong sexual conflict over allocation to sporophytes. Female gametophytes of pteridophytes produce a single sporophyte and often lack means of asexual reproduction. Therefore, sexual conflict is predicted to be attenuated. Finally, we explore similarities among models of mate choice, offspring choice and segregation distortion.     

Parthenogenetic female populations in the brown alga Scytosiphon lomentaria (Scytosiphonaceae,Ectocarpales): decay of a sexual trait and acquisition of asexual traits

In isogamous brown algae, the sexuality of populations needs to be tested by laboratory crossing experiments, as the sexes of gametophytes are morphologically indistinguishable. In some cases, gamete fusion is not observed and the precise reproductive mode of the populations is unknown. In the isogamous brown alga Scytosiphon lomentaria in Japan, both asexual (gamete fusion is unobservable) and sexual populations (gamete fusion is observable) have been reported. In order to elucidate the reproductive mode of asexual populations in this species, we used PCR‐based sex markers to investigate the sex ratio of three asexual and two sexual field populations. The markers indicated that the asexual populations consisted only of female individuals, whereas sexual populations are composed of both males and females. In culture, female gametes of most strains from asexual populations were able to fuse with male gametes; however, they had little to no detectable sexual pheromones, significantly larger cell sizes, and more rapid parthenogenetic development compared to female/male gametes from sexual populations. Investigations of sporophytic stages in the field indicated that alternation of gametophytic and parthenosporophytic stages occur in an asexual population. These results indicate that the S. lomentaria asexual populations are female populations that lack sexual reproduction and reproduce parthenogenetically. It is likely that females in the asexual populations have reduced a sexual trait (pheromone production) and have acquired asexual traits (larger gamete sizes and rapid parthenogenetic development).     

GAMETOPHYTE ONTOGENY,SEX EXPRESSION,AND GENETIC LOAD AS MEASURES OF POPULATION DIVERGENCE IN BLECHNUM SPICANT

Early gametophyte ontogeny was quantitatively distinct for Olympic Peninsula, Alaskan, and disjunct Idaho populations of the homosporous fern Blechnum spicant (L.) J. Sm. Although variable, gametophyte sex expression was shown to have a genetic component. Statistically different patterns of sex expression characterize each population. The Olympic Peninsula populations were distinct from each other but consistent in having a predominantly unisexual pattern. The disjunct Idaho population was predominantly bisexual at the time when comparable field collected gametophytes bear sporophytes. Preliminary experiments suggest that an antheridogen operates in this species. Increased sowing density favors maleness, and an extract from soil cultures of gametophytes shifts cultures to an exclusively male pattern after a dramatic suppression of growth. Mating experiments revealed that all populations are interfertile, although fertility was highest when the test Idaho population underwent intergametophytic-selfing. The Idaho population evidenced a low level of genetic load consistent with predictions based on its sex expression. Although Olympic Peninsula populations evidenced apparent high genetic load in some experiments, failure to produce abundant sporophytes in other experiments suggested that additional cultural factors operated to reduce sporophyte formation. Moderate density mating experiments produced single sporophytes that were comparable to field collections. Isolated gametophytes underwent polyembryony after a time delay and gametophyte proliferation. Cultural conditions which allow sporophyte formation on isolated gametophytes without this delay or proliferation must be sought before further genetic analysis is undertaken.     

不同环境因子对珍稀观赏蕨类金毛狗配子体性别分化的影响

金毛狗[Cibotium barometz(L.)J.Sm.]是珍稀观赏蕨类的重要类群,为国家二级重点保护野生植物。该研究以金毛狗孢子为试验材料,探究培养密度、外源赤霉素以及光质等不同环境因子对金毛狗配子体性别分化的影响,为金毛狗人工繁育和蕨类植物配子体性别决定机制研究提供技术支持。结果表明:(1)低配子体培养密度(1个/cm2和5个/cm2)有利于颈卵器和雌配子体形成,随着配子体培养密度增加,颈卵器平均数量及雌配子体比率下降,精子器平均数量以及雄配子体和两性配子体比例增加,但配子体培养密度过高(80个/cm2)会导致大量无性配子体产生。(2)不同配子体培养密度下,随着培养时间延长,两性配子体比率均有增加,且增加幅度基本一致。(3)外源GA4显著抑制颈卵器和雌配子体形成,并显著促进精子器和雄配子体形成;外源GA3对金毛狗配子体性别分化没有显著影响。(4)白光、红光、蓝光等不同光质对金毛狗配子体性别分化未产生显著影响,但会影响配子体的发育和形态建成。     

The reproductive biology of Polytrichum formosum: clonal structure and paternity revealed by microsatellites

Using highly polymorphic microsatellite markers, we assessed clonal structure and paternity in a population of the bryophyte species Polytrichum formosum. Identical multilocus genotypes of individual shoots were almost never observed in spatially separated cushions, but were found to be highly clustered within moss cushions. Therefore, asexual reproduction through dispersal of gametophyte fragments is not very important in P. formosum. However, asexual reproduction on a very localized scale through vegetative growth of genets (branching of gametophytes via clonal growth of rhizomes) is very extensive. The patchy spatial distribution of genets and the absence of intermingling among genets suggest that this species follows a 'phalanx' clonal growth strategy. Vegetative proliferation of genets will increase their size, and, consequently, will have considerable fitness consequences for individuals in terms of increased genet longevity and reproductive output. Although paternity analysis of sporophytes confirmed male genet size, i.e. gamete production, to be an important determinant of male reproductive fitness, it also showed that the spatial distance to female genets is the predominant factor that governs male reproductive success. Moreover, we showed that male gamete dispersal distances in P. formosum are much further than generally assumed, and are in the order of metres rather than centimetres. Combining the findings, we conclude that the high genotypic diversity observed for this facultatively clonal species is most likely explained by a preponderance of sexual reproduction over clonal reproduction.     

华南紫萁和粗齿紫萁的配子体发育研究

以腐叶土为基质 ,对华南紫萁和粗齿紫萁进行了孢子繁殖 ;利用Olympus显微镜观察和记录了它们的孢子萌发和配子体发育过程。结果表明 ,两者的孢子及配子体性状极为相似 :孢子同型 ,三裂缝 ,壁厚 ,含叶绿体 ,萌发快 ;在常温下 ,孢子的存活期短 ;孢子萌发需要光 ,萌发类型为紫萁型 ;丝状体阶段不发达 ,仅 2～ 3个细胞 ;原叶体无毛状体 ,中脉厚 ,精子器和颈卵器较大 ;由原叶体发育成幼孢子体所需时间极长。不同之处 :粗齿紫萁的孢子和精子器大些、颈卵器成对着生、成熟配子体多数为雌雄异株、受精率极低。华南紫萁和粗齿紫萁的孢子及配子体发育特点与紫萁、分株紫萁、绒紫萁、桂皮紫萁的极为相似 ,它们既具有大量的原始性状 ,也具有少数进化性状 ,支持把紫萁科列入原始薄囊蕨纲的观点。此外 ,也从孢子和配子体性状方面说明了紫萁属是一个自然的分类群     

What limits male range sizes at different population densities? Evidence from three populations of water voles

In many mammalian species, females compete with each other for food and space to raise offspring, while males compete with each other for access to females. Few studies have examined the factors which limit male range sizes or the degree of overlap between male ranges. We deduced four possible responses of the range -sizes of non-territorial male small mammals to increasing population density and/or levels of forage abundance. These were: (1) male range sizes might remain the same irrespective of population density; (2) at high population densities males may become territorial, and their ranges small and non-overlapping; (3) at high forage abundance/population densities, males' ranges may become smaller but remain intra-sexually overlapping; (4) at high forage abundance, male range sizes may increase. We examined the relationship between population density, range lengths and range overlaps and body weights of both sexes in a comparative study of three populations of water voles Arvicola terrestris . Male range sizes were smaller at higher population densities, but their ranges remained both inter- and intra-sexually overlapping. Heavier males had larger ranges than did lighter males at all sites. These results comply with what would be expected if male range sizes were at least partially restricted by the number of ranges of other individuals with which they overlapped. Although we could not discount the hypothesis that forage abundance may also have had a direct effect on male range sizes, our results implied that male range sizes were at least partially determined by social factors.     

Interference of asexual and sexual reproduction in the green hydra

The green hydra, Hydra viridissima, has three sexes: hermaphrodite, male, and female. I investigated the reproductive strategies of the green hydra and the relationship between asexual budding and sexual reproduction. The proportion of mature individuals in the asexually reproducing population increased with increasing temperature. Sexual reproduction did not interrupt asexual budding in hermaphrodites or males; sexual reproduction did interrupt asexual budding in females. Sexual reproduction also resulted in exponential population growth during the reproductive season. The number of asexual buds on each parental individual was positively correlated with the parental individual size in asexual individuals and in males. The same positive correlation was found between the number of testicles and the size of males. These correlations reflect a common tendency in asexual and sexual reproduction: larger parental individuals have a greater number of propagules or gametes. No correlation was found between size and buds or size and gonads in hermaphrodites; hermaphrodites had at most one asexual bud and were significantly larger than males, females, and asexual individuals. The larger size of hermaphrodites supports the hypothesis that producing both female and male gonads is more energetically costly than producing only one type of gamete (gonochorism).     

Towards an understanding of sex determination in Equisetum: an analysis of regeneration in gametophytes of the subgenus Equisetum

Equisetum gametophytes possess extensive powers of regeneration without showing any signs of diminished vigour or abnormal development. Isolated lamellae, pieces of antheridial and archegonial branches were used to analyse gametophyte morphogenesis and sex determination. Intact individuals and subcultured sexual branches never revert to a vegetative state. Transplantation of apical pieces of archegonial branches on to fresh medium delays the onset of antheridium formation indicating phenotypic control. On the other hand, male branches invariably continue producing antheridia. Experimentally induced adventitious branches on lamellae are interpreted as the result of physiological isolation from inhibitors. In intact individuals these adventitious branches are always antheridial: after subculturing, archegonial branches are also produced, demonstrating that the gametophytes of Equisetum are all potentially bisexual. Isolated lamellae regenerate secondary gametophytes, the sexual ratios between which vary between species, but lamellae from archegonial tissues invariably produce more female individuals, probably reflecting the different sizes of the respective lamellae. The view that mixtures of intact male and bisexual Equisetum gametophytes result from inherent physiological differences between the spores is not supported.     

不同光照度下水蕨配子体叶绿素荧光特性的研究

以模式植物水蕨的两性配子体和雄配子体为材料,研究了不同光照度下蕨类植物配子体叶绿素荧光参数的差异,探讨水蕨配子体光系统Ⅱ(PSⅡ)对不同光照的响应规律.结果表明:同一光照度条件下,水蕨两性配子体的实际光化学效率(Ф_(PSⅡ))和潜在光化学效率(F_v/F_m)均高于雄配子体,但无显著差异,此时两者间的光化学猝灭系数(q_p)和非光化学猝灭系数(q_N)值也无显著差异;在4 000 lx光照条件下,两性配子体PSⅡ驱动的电子传递速率(ETR_(PSⅡ))显著高于雄配子体.随着光照度的增加,水蕨两性配子体的Ф_(PSⅡ)、ETR_(PSⅡ)和q_p值逐渐增加,且在4 000 lx光照下显著增加,但F_v/F_m和q_N值无显著变化;而雄配子体的Ф_(PSⅡ)、ETR_(PSⅡ)、F_v/F_m、q_p和q_N值均呈先增后减趋势,但各光照处理间无显著差异.研究发现,水蕨两性配子体PSⅡ的活性相对比雄配子体高;两性配子体和雄配子体的光保护能力相似;光照度对水蕨雄配子体的光合能力没有显著影响,而4 000 lx的光照度对水蕨两性配子体的生长最有利.     

GROWTH AND DEVELOPMENT OF THE MALE GAMETOPHYTE OF GINKGO BILOBA WITHIN THE OVULE (IN VIVO)

The mature male gametophyte of Ginkgo biloba can be divided into two regions: a large saccate structure that is suspended within the fertilization chamber above the archegonia, and a pervasive, highly branched haustorial system that ramifies through the intercellular air spaces of the apex of the nucellus. This morphology appears to differ in many ways from the simpler more typical male gametophytes of most other groups of seed plants. Growth and development of the male gametophyte of Ginkgo biloba were studied using computer reconstruction techniques to generate images of the gametophyte from data derived from serial sections through the ovule. These investigations reveal that morphological development of the male gametophyte of Ginkgo biloba is divided into three distinct phases: 1) Germination, characterized by an initial brief period of diffuse growth. This phenomenon has not been described for any other seed plant male gametophyte; 2) Initiation of tip growth and the formation of a tubular body, as typifies all seed plant male gametophytes. In Ginkgo, this is accompanied by a high degree of branching, giving rise to an extensively branched haustorial system; 3) Late swelling of the proximal unbranched portion of the gametophyte resulting in formation of the saccate structure that is characteristic of the mature gametophyte. This process appears to be very similar to late development in cycad male gametophytes. Thus, despite the seemingly anomalous morphology of the mature male gametophyte of Ginkgo biloba, specific patterns of growth and development are in many ways similar to growth processes expressed by the male gametophytes of some or all major groups of seed plants.     

INTERPOPULATIONAL VARIATION IN THE GAMETOPHYTES OF PELLAEA ANDROMEDAEFOLIA

The development and mature morphology of the gametophytes from both sexual and apogamous populations of the fern Pellaea andromedaefolia were investigated. While most sexual examples were indistinguishable, some differences were noted. An insular collection was distinctive in its variability and irregularity of form. Although the latter was a representative of var. pubescens, other collections of the variety could not be distinguished from var. andromedaefolia on the basis of gametophytic characteristics. The apogamous gametophytes were decidedly more variable in development and often very different from sexual thalli. The mature asexual thalli tended to be more irregular in form and usually sharply divergent from the typical cordate type characteristic of the sexual populations. Each of the five apogamous samples was unique with respect to gametophyte development. The differences among the gametophytes of the various populations do not correlate with the sporophytic characteristics which differentiate the two varieties of the species.     

Density-dependent sexual selection in external fertilizers: variances in male and female fertilization success along the continuum from sperm limitation to sexual conflict in the sea urchin Strongylocentrotus franciscanus

Sperm competition and female choice are fundamentally driven by gender differences in investment per offspring and are often manifested as differences in variance in reproductive success: males compete and have high variance; most females are mated and have low variance. In marine organisms that broadcast spawn, however, females may encounter either sperm limitation or sperm competition. I measured the fertilization success of male and female Strongylocentrotus franciscanus over a range of population densities using microsatellite markers. Female fertilization success first increased and then decreased with mate density, limited at low density by sperm limitation and at high density by polyspermy. Mate density affected variance in fertilization success in both males and females. In males, the variance in fertilization success increased with mate density. In females, the pattern was more complex. The variance in female success increased similarly to males with increased mate density but then decreased to low levels at intermediate densities, where almost all eggs were fertilized. As density increased further, the female variances again increased as polyspermy lowered average fertilization success. Male and female variances differed only at intermediate densities. At low densities, both sexes may be under selection to increase fertilization success; at intermediate densities, males may compete; and at high densities, both sexes may be under selection to increase success by increasing (males) or decreasing (females) likelihood of fertilization during sexual conflict. Only within a narrow range of densities do patterns of sexual selection mirror those typically noted in internally fertilizing taxa.     

Population biology of the rare copper moss,Scopelophila cataractae

The so-called copper mosses include a number of rare species that exhibit very broad intercontinental geographic distributions comprised of highly disjunct occurrences. In one species, Scopelophila cataractae, only the haploid gametophyte generation exists in the United States, although sporophytes occur in tropical America and in Asia. Gametophytic plants were sampled from all U.S. populations to determine what factors limit sexual reproduction. More than 50% of the plants in every population were devoid of gametangia, and no population contained plants with both male and female gametangia. Morphological differences between plants from putative male and female populations (when each was interpreted to be unisexual) suggested gametophytic sexual dimorphism, but generalized sexual differences were not maintained under common garden conditions. Experimental growth of plants on soils with low, moderate, or high concentrations of metals demonstrated extensive morphological variability, and thus genetic polymorphism, among five asexual populations. Morphological traits were also significantly plastic in response to differing soil types, but there was no evidence of differences in patterns of plasticity between sexes or populations. All populations produced higher cover area and individual plants formed larger leaves on the most highly metal-contaminated soil. Populations varied significantly in growth on less contaminated soil, again suggesting genetic variability.