被子植物雌全同株性系统:系统演化、性表达与进化意义

雌全同株是指雌花和两性花共同发生在同一植株上的性表达形式。作为被子植物从雌雄同花(两性花)向雌雄同株异花进化的一个重要阶段,雌全同株性系统在减少昆虫对雌性的取食和伤害、提高异交率以减少近交衰退、减少雌/雄功能干扰、提高雌/雄性功能间资源分配的灵活性,以及吸引传粉者等方面具有重要的进化适应意义。根据APG III分类系统,雌全同株性系统在被子植物木兰分支(magnoliids)的短蕊花科、单子叶植物分支(monocots)的天南星科和禾本科,以及核心真双子叶植物分支(core eudicots)中的菊科、苋科、唇形科和石竹科等23科中均有报道,且以菊科植物中最多。雌全同株植物不同类群的雌花和两性花在位置、形态、大小及开花时间等性表达特征上表现出多样化,且这些特征不仅受遗传因子的调控,还受可获得资源(如营养、光照、温度和水分等条件)的制约。该文针对我国对雌全同株性系统的研究还相对较少的现状,重点对具雌全同株性系统的类群在被子植物中的分布与系统演化、性表达与环境的关系等方面进行了分析与总结,并对有关其进化适应意义的5个假说进行了介绍和评价,对今后的研究方向进行了展望,以期为推动我国对被子植物雌全同株性系统的进化式样与机制研究提供理论资料。     

七叶树繁育系统的研究

对七叶树的开花物候期、花部特征、单花开放动态、花粉活力、柱头可授性、花粉-胚珠比(P/O)和杂交指数(OCI)等有关生殖生物学特性进行了研究。结果表明:(1)在西安地区七叶树4月22日进入初花期,5月5日至13日为盛花期,5月14日进入终花期,整个群花期持续30d左右,单花花期持续5～8d。(2)七叶树的花有雄花和两性花两种形态,其在形态大小和生理上有显著差异,雄花小于两性花,其子房退化,雄花的花粉活力远远大于两性花的花粉活力,雄花开花当天花粉活力最强,达到75.69%,是七叶树授粉的最佳时间;两性花雌雄蕊发育正常,其柱头可授期持续8～9d,开花后3d柱头可授性达到最强。(3)P/O和OCI检测结果均表明,七叶树繁育系统为异交,需要传粉者,属于雄全同株植物。     

多伞阿魏的繁育系统及其生态适应性

多伞阿魏（Ferula ferulaeoides）是新疆特有的类短命植物,具有巨大的生态作用和潜在的经济价值。实验通过研究多伞阿魏的花部形态特征、繁殖系统及传粉昆虫,来阐明其对生态环境的响应。结果表明：（1）多伞阿魏花序花期一般持续10~15 d,单花花期为7~10 d,两性花呈现雌雄异位和雌雄异熟的现象;（2）其繁育系统类型为兼性异交,存在自交部分亲和,自然结果率较高,为72.24%;（3）传粉昆虫多种多样,其中以双翅目和膜翅目昆虫居多;（4）开花当天的花粉活力呈单峰曲线,在12：00时活力最高,柱头可授性在花药完全散粉后第2天最强。     

雄全异株植物瘿椒树（省沽油科）的传粉生物学

从开花动态、传粉昆虫、花的形态结构、繁育系统、花粉活力和柱头可授性等方面研究了我国特有珍稀植物瘿椒树（Tapiscia sinensis Oliv.）的传粉生物学特性。瘿椒树是典型的雄全异株植物,两性花中含有功能性花粉,且自交亲和,但雄花花粉活力和萌发力是两性花的10倍以上。雄株和两性植株具有相同开花物候期,花期均为5月下旬至6月上旬,单花期为4-5天,雄花和两性花的5枚花药开裂的不同步性明显延长了散粉时间。两性花雌蕊先熟,柱头可授性较长。具有适应风媒和虫媒传粉的花部特征。传粉昆虫主要为蜜蜂科（Apidae）和食蚜蝇科（Syrphidae）昆虫,访花高峰期为8：30-10：30。维持瘿椒树雄全异株的可能机制是：雄株总体上增加了异交花粉的数量和质量;两性花的雄蕊为该物种提供了繁殖保障,同时为传粉者提供了报酬。     

新疆阿魏雄性不育的细胞形态学研究

新疆阿魏是特产于我国新疆的伞形科阿魏属多年生一次结实草本植物,属于国家二级保护的濒危植物。其种群中除了该科植物典型的雄全同株个体以外,还具有一定数量雄性不育的雌株。为了探究新疆阿魏的雄性不育现象及其影响因素,该文从细胞形态学角度对种群中的雌株以及雌花的形态特征进行了观测,采用石蜡切片技术对功能性雌花雄蕊的花药败育过程进行了观察。结果表明：(1)雌株3月底萌动,4月中旬进入花期,5月底果实成熟;物候期与雄全同株个体相同。(2)植株高度(71.00±10.92) cm和直径(71.67±17.64) cm、一级分支(23.83±2.04);基生叶长(33.41±11.63) cm、宽(24.47±8.60) cm;在植株大小、基生叶大小等方面雌株与雄全同株个体无差异。(3)在雌株上,一级分支和二级分支均为雌花序,均可结实;雌花序的伞幅数/每复伞花序(13.22±4.70)、花数/每花序(12.03±2.30)、总花数/每复伞花序(159.08)均高于两性花序;雌株比雄全同株个体产生更多可结实的花,形成更多具有杂种优势的异交后代。(4)雌花序中,花排列紧密,花间距小于两性花序和雄花序;开花时...     

雄全异株植物瘿椒树(省沽油科)的传粉生物学

从开花动态、传粉昆虫、花的形态结构、繁育系统、花粉活力和柱头可授性等方面研究了我国特有珍稀植物瘿椒树(Tapiscia sinensis Oliv.)的传粉生物学特性。瘿椒树是典型的雄全异株植物,两性花中含有功能性花粉,且自交亲和,但雄花花粉活力和萌发力是两性花的10倍以上。雄株和两性植株具有相同开花物候期,花期均为5月下旬至6月上旬,单花期为4-5天,雄花和两性花的5枚花药开裂的不同步性明显延长了散粉时间。两性花雌蕊先熟,柱头可授性较长。具有适应风媒和虫媒传粉的花部特征。传粉昆虫主要为蜜蜂科(Apidae)和食蚜蝇科(Syrphidae)昆虫,访花高峰期为8:30-10:30。维持瘿椒树雄全异株的可能机制是:雄株总体上增加了异交花粉的数量和质量;两性花的雄蕊为该物种提供了繁殖保障,同时为传粉者提供了报酬。     

矮慈姑的传粉机制与交配系统

对矮慈姑（ＳａｇｉｔａｒｉａｐｙｇｍａｅａＭｉｑ）的传粉过程与花粉流作了观察,用同工酶遗传标记法对其一个自然居群的异交率作出定量估计。在自然及人工居群中均观察到虫媒传粉,其中有较大比例的近距离传粉。未检出风媒花粉流。自交可育,同时存在雌雄同株和花序内雌雄花异熟等异交机制。异交率估计结果（ｔ＝４９９％）表明其交配系统为异交／自交兼性系统。作者认为,其较大比例的自交应归因于同株之不同克隆分株间交配的频繁发生     

珍稀濒危植物距瓣尾囊草(Urophysa rochkii)的开花物候和繁育系统特性

通过野外观察,运用套袋授粉和联苯胺-过氧化氢法等实验方法对距瓣尾囊草开花物候以及种群的繁育系统特性进行研究。研究结果:1)距瓣尾囊草为两性花,聚伞状花序,花期较长,从头年12月持续到次年4月,单花花期持续8—15 d,种群花期可持续4个月左右。2)距瓣尾囊草在花朵未开放前花粉没有活力,花药开裂当天花粉活力达到94.8%,第2天为90.3%,第3天为81.6%,第4天为62.9%,此后其花粉活力明显减弱;柱头可授性于散粉后第3天开始出现,散粉后4—5天最强,散粉后第8天柱头失去可授性;柱头可授性和花粉活力有5 d左右的重叠期。3)其花粉胚珠比P/O为938.5±250,按照Cruden标准划分,其有性繁育系统为兼性异交。4)按照Dafni的标准,距瓣尾囊草的OCI=4,即繁育系统为异交、部分自交亲和、需要传粉者。5)人工异花授粉结实率达80%以上,略高于自然结实率78.94%;直接套袋结实率为5.71%;去雄套袋和去雌套袋均不结实。以上结果表明,距瓣尾囊草繁育系统表现为异交为主、部分自交亲和并且不存在无融合生殖现象,其开花物候、花部特征和繁育系统为适应特殊的环境提供了一定的生殖保障,本次研究结果为距瓣尾囊草的迁地保护和种群的繁衍复壮提供理论依据。     

两种姜科花柱卷曲性植物柱头的位置与其可授性的关系

花柱卷曲性是一种见于姜科山姜属(Alpinia)和砂仁属(Amomum)植物中的独特的性二态现象, 具有这一性系统的植物, 其居群包括上举型和下垂型两种表型, 所有个体的雌雄性别功能在时间和空间上分离。本研究通过操控授粉和花粉管生长两项实验, 探讨花柱卷曲性植物个体两性功能的隔离方式及其适应意义。云南草蔻(Alpinia blepharocalyx)操控授粉实验表明, 自花花粉对其异交率影响不显著(P > 0.05), 但可能由于自花花粉沉降导致胚珠贴现进而引起近交衰退, 操控条件下每果结籽数显著减少(P < 0.01)。云南草蔻和红豆蔻(Alpinia galanga)花粉管生长实验都显示无论对两种表型进行怎样的授粉处理, 花粉粒萌发和花粉管生长的速率仅在柱头处于可授位置时表现出最大值, 与自交和异交授粉方式无关; 上举型植株上午花粉囊虽未裂但其内花粉已成熟。研究结果表明即使不考虑花柱运动, 山姜属植物也具有异型雌雄异熟的特性。这一结果证实了花柱卷曲运动机制是通过互补式雌雄异位和异型雌雄异熟相结合形成的花部二态性, 异型雌雄异熟促进了异交, 而花柱运动的功能可能在于避免雌雄干扰。     

论雌花两性花同株植物的适应意义

雌花两性花同株是植物个体上同时有雌花和两性花的性系统。与其他性系统相比, 雌花两性花同株一直较少受到关注。尽管只有约3%的被子植物是雌花两性花同株的物种, 但其分布广泛, 具有多种多样的花部特征和传粉模式。解释雌花两性花同株物种的适应意义已有多种假说, 包括促进异交、避免雌雄干扰、优化资源配置、昆虫的植食作用以及增大对传粉者的吸引力等, 但证据不多, 并且研究对象主要集中在菊科Asteraceae。进一步对不同类群、不同传粉模式的雌花两性花同株的物种进行繁殖生物学研究, 并结合系统发育的比较研究, 有助于深入理解这一性系统的起源与进化。     

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     

Sex Expression and the Evolutionary Advantages of Male Flowers in an Andromonoecious Species, Sagittaria guyanensis subsp. lappula (Alismataceae)

Sex expression in Sagittaria guyanensis H.B.K. subsp. lappula (D. Don) Bojin is typically andromonoecious while the other species in the same genus are basically monoecious. The evolutionary advantages of male flowers and hermaphrodite flowers in S. guyanensis subsp. lappula were assessed by measuring sex allocation and pollen movements in two wild populations of the species. Two cultivated populations served as controls. The percentage of male flowers was very low in the two wild populations in Dongxiang, Jiangxi Province and Wuyishan, Fujian Province, viz., 2.48% and 0.96% respectively. In the two cultivated populations, male flower percentage significantly increased when the soil was of higher nutrient content. This indicates that the allocation to male versus female reproduction might change in response to environmental factors. Pollen production per male flower was 4.1 times higher than that of a hermaphrodite flower. The floral shape and size of male and hermaphrodite flowers were similar. No difference was observed between these dimorphological flowers in pollen germination rate in vitro and in the speed of pollen tube growth in vivo. Anthesis was only 4-5 h. Male flowers usually opened 0.5 h earlier than hermaphrodite flowers. An unexpected finding was that no pollen from the male flowers was found on the stigmas of the hermaphrodite flowers, in spite of the occasional visits by insects to both types of flowers in both wild and control populations. A consistent pattern of fruit development was found to exist in open pollinated flowers as well as in flowers that had been bagged. The sex ratios did not have significant influence on fruit set. Approximately 25% of the pistils in a gynoecium failed to develop into fruits because no pollen was deposited on them, indicating that the fruit set of this andromonoecious plant is mainly affected by pollen limitation rather than resource limitation. Reproduction in S. guyanensis subsp. lappula in the habitats was dependent on self-pollination in hermaphrodite flowers. The male flowers in this species might be a potential source of additional pollination and may facilitate cross-pollination. The fact that the flowers of monoecious species in Sagittaria pollinated by a wide diversity of insect visitors may contribute enormously to the diversification of sex expression in this genus.     

SEXUAL ALLOCATION AND ANDROMONOECY: RESOURCE INVESTMENT IN MALE AND HERMAPHRODITE FLOWERS OF SOLANUM CAROLINENSE (SOLANACEAE)

Male and hermaphrodite flowers from the andromonoecious perennial Solanum carolinense were compared. Hermaphrodite flowers were 11–16% heavier than males, and this difference was consistent for two clones studied. Hermaphrodite flowers contained a greater amount of nitrogen than did male flowers, but males had higher percent nitrogen. Male and hermaphrodite flowers had equal investment in male structures: stamen mass, number of pollen grains, mass of pollen grains ejected by vibration, and pollen size were all similar for the two flower types. In contrast, male flowers had reduced masses of disfunctional female structures: ovaries of males were ⅓ as massive, and styles 1/7; as massive, as those of hermaphrodites. These results are consistent with the hypothesis that male flowers (which occur where fruit set is unlikely) increase male fitness through pollen donation, while saving resources that would otherwise be invested in hermaphrodite flowers that abscise.     

雄全同株植物簇花芹花期性别分配与开花式样

簇花芹(Soranthus meyeri)是古尔班通古特沙漠中常见的、具雄全同株性系统的伞形科多年生早春短命植物。该文对簇花芹花期性比(两性花数/总花数)与植株大小的关系及其开花式样进行了研究, 重点对花期大小依赖的性别资源分配进行了讨论。结果表明: 2006-2008年簇花芹群体水平的性比分别为0.69 ± 0.03、0.62 ± 0.03和0.69 ± 0.02, 彼此间无显著差异( p > 0.05), 表明其性比是相对稳定的, 可能受遗传因素的控制。雄花生物量与花粉量均比两性花的小, 说明产生雄花比产生两性花所需资源少。一级复伞形花序比二级复伞形花序具有较多的两性花, 说明前者易从植株上获得资源用于增加雌性适合度; 而后者产生较多的雄花以避免在雌性功能上资源投入的浪费, 增大花展示以吸引更多传粉者来增加花粉输出总量, 提高其整体适合度。植株水平的性比与地上营养器官的生物量间呈正相关关系, 说明较大个体对雌性功能的投资较大, 雌性繁殖成功受资源限制。复伞形花序内各伞形花序几乎同时向心开放, 且所有两性花及花序均为雄性先熟, 雌雄阶段完全分离, 但一级复伞形花序比二级复伞形花序早开放约5天, 彼此开花重叠期约为1天。这些特征对于一级复伞形花序进行异株异花授粉以及植株内不同级别花序间的同株异花传粉、避免雌雄功能间的干扰具有重要意义。     

Subdioecy in Consolea spinosissima (Cactaceae): breeding system and embryological studies

The breeding system and the embryology of Consolea spinosissima, a tree-like opuntioid endemic to Jamaica, were investigated. Morphological and embryological studies revealed that the species is subdioecious, with three sexual morphs present in the 150 × 120 m plot studied at Hellshire Hills, Jamaica. The female morph has pistillate flowers with open stigma lobes, no pollen grains, and sets fruit. The male morph has cryptic staminate flowers with closed stigma lobes, viable pollen grains, and a nonfunctional gynoecium that does not set seed. The weak hermaphrodite morph has low fruit set and "perfect" flowers that superficially resemble the functionally staminate flowers of the male morph. These perfect flowers reach anthesis with viable pollen grains, with no or only a few functional ovules, and with the style supporting pollen tube growth. Embryological studies showed that the critical stage for sex determination occurs earlier in pistillate than in staminate and perfect flowers. Anthers of pistillate flowers abort prior to microspore tetrad formation, whereas ovules of the staminate and perfect flowers degenerate after the complete maturation of the embryo sac. Based on flower structure and embryological data, we hypothesize that the ancestor of C. spinosissima is/was hermaphroditic.     

Effects of pollen reward removal on fecundity in a self-incompatible hermaphrodite plant

The pollen of hermaphrodite plants is often utilised by flower-visiting animals. While pollen production has obvious benefits for plant male fitness, its consequences for plant female fitness, especially in self-incompatible hermaphrodite species, are less certain. Pollen production could either enhance seed production though increased pollinator attraction, or reduce it if ovules are discounted by deposition of self pollen, as can occur in species with late-acting self-incompatibility. To test the effects of pollen reward provision on female fitness, we artificially emasculated flowers in two populations of the succulent Aloe maculata (Asphodelaceae), which has a late-acting self-incompatibility system, over the course of its flowering period. Flowers of this species are visited by sunbirds (for nectar) and native bees (for pollen and nectar). We measured floral visitation rates, floral rejection rates, pollen deposition on stigmas and fruit and seed set in both emasculated and non-emasculated plants. We found that flowers of emasculated plants suffered reduced visitation and increased rejection (arrival without visitation) by bees, but not by sunbirds; had fewer pollen grains deposited on stigmas and showed an overall decrease in fruit set and seed set. Rates of seed abortion were, however, greatly reduced in emasculated flowers. This study shows that pollen rewards can be important for seed set, even in self-incompatible plants, which have been assumed to rely on nectar rewards for pollinator attraction. Seed abortion was, however, increased by pollen production, a result that highlights the complexity of selection on pollen production in hermaphrodite flowers.     

Yield improvement through female homosexual hybrids and sex genetics of sweet gourd (<Emphasis Type="Italic">Momordica cochinchinensis</Emphasis> Spreng.)

The present research aimed to evaluate the effect of silver nitrate (AgNO₃) on sex modification in sweet gourd (Momordica cochinchinensis Spreng.), and to explore the possibility of sexual crossing between two genetically female plants. Spray applications of AgNO₃ on 30 days’ old female plants induced hermaphrodite flowers. Male plants were insensitive to the AgNO₃ sprays. Application of 500 mg/l AgNO₃ on female plants produced the maximum proportion of induced hermaphrodite flowers. Hermaphrodite flowers appeared 17–21 days after AgNO₃ spray and continued up to 8–17 days, depending upon the concentration of AgNO₃. Pollen grain viability of induced hermaphrodite flowers (93.5%) was similar to pollen grain viability of normal male (95.0%) flowers. Because of higher fruit weight, progenies from female homosexual cross recorded higher yield. The hybrids from such crosses produced only female plants while the hybrids between female and normal male segregated into male and female in an equal proportion, indicating that sex in sweet gourd is controlled by a single factor, male being heterozygous and female being homozygous recessive. Through this technique, elite characters of female genotypes could be combined into a single plant.     

Reproductive characters in a gynodioecious species, Silene italica (Caryophyllaceae), with attention to the gynomonoecious phenotype

Gynodioecious populations (i.e. populations with female and hermaphrodite individuals) often contain a third phenotype with an intermediate sex expression. In Silene italica , this phenotype is characterized by a mixture of pistillate and perfect flowers and is thus gynomonoecious. To characterize sexual functions of these gynomonoecious individuals and their potential influence in the maintenance of gynodioecy, reproductive characters of the three sexual phenotypes were compared over 2 years in several families of S. italica produced by crossing. We found that gynomonoecious individuals were intermediate for flower production and female fertility characters, although they did not always significantly differ from female individuals. Perfect flowers of gynomonoecious and hermaphrodite plants were similar in size and pollen production. Gynomonoecious individuals were thus intermediate in female and in male functions. Family effects were found for most of the characters. The female advantage (i.e. the fertility of females compared to the female fertility of pollen producing plants) was not dramatically different when gynomonoecious plants were taken into account.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 583–591.     

Maytenus obtusifolia Mart. (Celastraceae): a tropical woody species in a transitional evolutionary stage of the gynodioecy–dioecy pathway

‘Gynodioecy–dioecy’ is one of the pathways by which dioecy can arise from hermaphroditism. Studies on sex determination and development of gynodioecious systems have focused on temperate and/or annual species. Little is known about the evolutionary dynamics of gynodioecy and dioecy in perennial tropical species, where these systems have more frequently evolved. Maytenus obtusifolia is an abundant species in restingas in southeastern Brazil. The sexual system of M. obtusifolia was investigated by studying the floral structure and reproductive biology. We considered the sexual system as an intermediate step in the pathway gynodioecy-dioecy. The characterization of the hermaphrodite morph was complex, because of a gradient of variation in floral morphology and reproductive characteristics (sizes of the style and stigmatic surface, pollen viability, embryo sacs containing hypertrophied synergids, and fruit set). This variation leads to different proportions of functional male and female flowers among hermaphrodite plants and is responsible for the different levels of reproductive success. Female reproductive success and pollen viability were negatively correlated with the hermaphrodite morph (r = ?0.67). The higher fruiting intensity and fruit/flower ratio of females (41 %) compared to hermaphrodites (2 %) and the fact that female plants produce more and better-quality seeds support the female compensation. We suggest that female sterility may be linked to the set of changes in the carpels. The differences in the quality and quantity of pollen grains of hermaphrodite plants, and the similar individual rates of pollen viability observed for three consecutive flowering events, may indicate a relationship with nuclear cytoplasmic sex determination.     

Fruit set increases with maleness in the andromonoecious Acacia caven

In long-lived plants, sexual expression can be highly variable in time and space. The extent to which sexual dimorphism of flowers is expressed in the same or in separate individuals, may have implications for the plants’ dependence on pollen vectors, the compatibility of the pollen received, and the potential for setting fruit. Here, we account for variability in sexual expression in the small tree Acacia caven, which produces hermaphrodite and male flowers on the same or in separate individuals. We focus on whether individual sexual expression influences fruit set, which latter has been reported to be extremely low in this species. Individual sexual expression, estimated as the hermaphrodite: male flowers ratio was variable both within and among plants across two reproductive seasons. Though male flower production could be very high (up to 99% of the flowers), all trees produced both flower types within the same individual, thus confirming andromonoecy in this species. More interestingly, hermaphrodite: male flowers ratio had a negative effect on fruit set. This pattern was consistent across two reproductive seasons and suggests that maleness should favor female function. Similar trends found in other plant species with the same sexual system support the hypothesis that male flowers of andromonoecious species may contribute to reduce self-pollen interference compared with hermaphrodite flowers. By favoring the deposition of compatible pollen grains and increasing fruit set, maleness would then help to maintain andromonoecy in this and other andromonoecious species.