SEX EXPRESSION,BREEDING SYSTEM,AND POLLEN BIOLOGY OF RICINOCARPOS PINIFOLIUS: A CASE OF ANDRODIOECY?

A population of 54 Ricinocarpos pinifolius (Euphorbiaceae) plants contained male plants, which produced only staminate flowers, and hermaphrodites, which produced staminate and pistillate flowers. The fraction of pistillate flowers ranged continuously from 0 to 0.68. Insect pollination was effective and fruit set virtually complete except for losses to herbivores. Self pollen, outcross pollen from male plants, and outcross pollen from hermaphrodites were all equivalent in viability, germination, tube growth, ovule penetration, and fruit setting ability. Inbreeding depression was manifested as late abortion of some selfed seeds. Geitonogamous selfing is largely prevented by temporal separation of male and female functions within plants. This temporal separation, combined with population-wide synchrony of flowering, may create unusual conditions allowing male plants at low frequency to match hermaphrodites in reproductive success.     

POLLINATION ECOLOGY OF A SEAGRASS,THALASSIA TESTUDINUM (HYDROCHARITACEAE), IN ST. CROIX

The reproductive ecology of a dioecious Caribbean seagrass, Thalassia testudinum was studied in St. Croix, U.S. Virgin Islands. Flowering in Thalassia coincided with spring tides with anthesis and anther dehiscence in staminate plants occurring at night. As predicted by search theory, pollen is dispersed in negatively buoyant rafts of pollen grains (that superficially resemble strings of frog spawn) which are bound by a slime of apparent thecal origin. Dispersal of pollen is submarine and approximately two-dimensional along the plane of the substratum. The stigmas on pistillate plants are linear, stiff, and densely papillate; they protrude from the mouth of the subtending leaf. Floral ratios of staminate to pistillate flowers were 60 to 1 with an average distance of 4.0 cm from pistillate flower to the closest staminate flowers. Populations included a high proportion of short shoots with numerous (> 100) leaf scars indicating ages for these axes of up to 3–4 years. Nonflowering shoots can be sexed and frequency of flowering can be assessed from scars of old inflorescences. Preliminary results suggest that there is no regular pattern of flowering, since the interval between flowering events is variable.     

Pollen movement to flowering canopies of pistillate individuals of three rain forest tree species in tropical Australia

This study measured the quantities of effective pollen vectors and their pollen loads arriving at the canopies of dioecious tropical rain forest trees in north-east Queensland. Population flowering synchrony, effective pollinator populations and pollen loads transferred between staminate and pistillate trees were compared among three insect-pollinated tree species. All three were visited by a wide range of insects, 75% of which (mostly 3–6 mm long) carried conspecific pollen. Fewer than 8% of individual insects were found to be carrying single-species pollen exclusively and none could be described as specialist pollen foragers. The introduced honeybee carried greater quantities of pollen than any native species but was not necessarily a reliable pollinator. The brief flowering periods in Neolitsea dealbata (3–4 weeks) and Litsea leefeana (4–5 weeks) populations were synchronized among individuals. Flowering in the Diospyros pentamera population extended over 15 weeks and most individuals were in flower for most of this period. Staminate trees began flowering earlier, produced more flowers and attracted relatively more insects than did pistillate trees, suggesting a density-dependent response of pollinators to flowering performance. Pollen was trapped in greater quantities on insects at staminate trees than at pistillate trees. Insect numbers increased at peak flowering periods and Diptera were the most abundant flower visitors. Anthophilous Coleoptera were more numerous at staminate than at pistillate trees in all three tree species populations. Larger quantities of pollen were mobilized during peak flowering times although the greatest quantities were transferred to pistillate canopies towards the end of the population flowering periods. Diptera carried pollen more often to pistillate N. dealbata and L. leefeana trees than did other groups whereas Coleoptera carried pollen more often to pistillate D. pentamera trees. The two contrasting flowering performances in the three tree species are discussed with reference to mechanisms that facilitate pollen transfer between staminate and pistillate trees.     

Induction of male flowers in a pistillate line ofRicinus communis L. by silver and cobalt ions

Aqueous solutions of silver nitrate (10–100 g/plant) and cobalt chloride (125–500 g/plant), injected into the main stem of plants of the pistillate cv. 240 ofRicinus communis when the vegetative shoot apex was beginning to become reproductive, induced the formation of staminate (male) flowers with viable pollen in the normally strictly pistillate (female) terminal inflorescence, their number increasing with the dose of Ag⁺ and Co²⁺. No formation of bisexual flowers was noted. Female flowers pollinated with pollen from the induced male ones produced fruits and viable seeds.     

POLLEN REMOVAL AND POLLEN DEPOSITION AFFECT THE DURATION OF THE STAMINATE AND PISTILLATE PHASES IN CAMPANULA RAPUNCULOIDES

We examined factors affecting the duration of the staminate and pistillate phases in the protandrous flowers of Campanula rapunculoides L. (Campanulaceae). Under conditions of natural pollinator visitation, flowers experiencing low rates of pollen removal lasted significantly longer than flowers that had faster rates of pollen removal. Experimental manipulations showed that low levels of pollen removal resulted in extension of the staminate phase. Hand-pollinations in which we varied the amount and source of pollen showed that when the number of fertilized ovules within an ovary is low, senescence of the flower is delayed, resulting in extension of the pistillate phase. We also report on pollinator foraging patterns within the vertical inflorescences of C. rapunculoides and the limiting factor for seed set in this population. The results are relevent to recent suggestions that floral characters often serve to reduce interference between the sexual functions in cosexual plants.     

间伐强度对黄桥林区辽东栎生殖构件的影响

为了阐明抚育间伐强度对辽东栎生殖构件的影响,以黄土高原南部黄龙和桥山林区辽东栎中龄林为研究对象,对不同间伐强度(对照、10%、20%、30%)下近自然经营抚育间伐5年后的辽东栎平均单株个体大小、生殖构件产量、比率及空间分布格局特征等进行研究.结果表明: 间伐强度对辽东栎林分平均树高、胸径影响不显著,而与冠幅面积、冠幅体积呈显著正相关;辽东栎标准株的生殖枝、幼果、成熟果实产量随间伐强度的增大而增大,而营养枝、雄花序、雄花、雌花的产量变化不大.当间伐强度达到20%时,生殖枝、幼果、成熟果实产量显著提高,30%间伐下达到最大.生殖枝比例和座果率随间伐强度增大而增大,在20%、30%间伐强度时与对照的差异达到显著水平.约3/4的生殖枝、雌花、雄花集中分布于树冠上层,1/4分布于下层;树冠阳面生殖构件数量占65%,树冠阴面占35%.树冠下层、阴面生殖枝、雌花、雄花序的空间分布比例随着间伐强度的增大而增大;树冠上层、阳面生殖枝、雌花、雄花序分布比例随间伐强度的增大呈递减趋势.因此,选择30%间伐强度(保留郁闭度0.6)最佳,可以大幅提高辽东栎的座果率和成熟率,有效提高结实量和品质,为辽东栎林的采种及天然更新提供保证.     

The Control of Apical Bud Growth and Senescence by Auxin and Gibberellin in Genetic Lines of Peas

Pea (Pisum sativum L.) lines G2 (dwarf) and NGB1769 (tall) (Sn Hr) produce flowers and fruit under long (LD) or short (SD) days, but senesce only under LD. Endogenous gibberellin (GA) levels were inversely correlated with photoperiod (over 9-18 h) and senescence: GA20 was 3-fold and GA1 was 10- to 11-fold higher in flowering SD G2 shoots, and the vegetative tissues within the SD apical bud contained 4-fold higher levels of GA20, as compared with the LD tissues. Prefloral G2 plants under both photoperiods had GA1 and GA20 levels similar to the flowering plants under LD. Levels of indole-3-acetic acid (IAA) were similar in G2 shoots in LD or SD; SD apical bud vegetative tissues had a slightly higher IAA content. Young floral buds from LD plants had twice as much IAA as under SD. In NGB1769 shoots GA1 decreased after flower initiation only under LD, which correlated with the decreased growth potential. We suggest that the higher GA1 content of G2 and NGB1769 plants under SD conditions is responsible for the extended vegetative growth and continued meristematic activity in the shoot apex. This and the increased IAA level of LD floral buds may play a role in the regulation of nutrient partitioning, since more photosynthate partitions of reproductive tissue under LD conditions, and the rate of reproductive development in LD peas is faster than under SD.     

Anther structure and pollen development in Melicoccus lepidopetalus (Sapindaceae): An evolutionary approach to dioecy in the family

Anther and pollen development in staminate and pistillate flowers of dioecious Melicoccus lepidopetalus (Sapindaceae) were examined by light and electron microscopy. Young anthers are similar in both types of flowers; they consist of epidermis, endothecium, two to four middle layers and a secretory tapetum. The microspore tetrads are tetrahedral. The mature anther in staminate flowers presents compressed epidermal cells and endothecium cells with fibrillar thickenings. A single locule is formed in the theca by dissolution of the septum and pollen grains are shed at two-celled stage. The mature anthers of pistillate flowers differ anatomically from those of staminate flowers. The epidermis is not compressed, the endothecium does not develop fibrillar thickenings, middle layers and tapetum are generally persisting, and the stomium is nonfunctional. Microspore degeneration begins after meiosis of microspore mother cells. At anthesis, uninucleate microspores and pollen grains with vegetative and generative nuclei with no cytokinesis are observed. Some pollen walls display an abnormal exine deposition, whereas others show a well formed exine, although both are devoid of intine. These results suggest that in the evolution towards unisexuality, the developmental differences of anther wall tissues and pollen grains between pistillate and staminate flowers might become more pronounced in a derived condition, such as dioecy.     

罗汉果雄株不足、花期不遇研究

<正> 罗汉果为雌雄异株植物,栽培品种均需人工授粉才能结实。当前,区内外罗汉果生产上,普遍存在着雄株不足,花期不遇的问题,新区更为严重,给生产带来严重的损失,成为目前急待解决的问题。 1981年以来,在科委、外贸,供销等部门大力支持下,我们通过新老产区雄株生长情况、     

MORPHOLOGY,VASCULAR ANATOMY AND EMBRYOLOGY OF PISTILLATE AND STAMINATE FLOWERS OF ASPARAGUS OFFICINALIS

Flowers of three pistillate (female), two heterogametic staminate (male) and two homogametic male genotypes of Asparagus officinalis L. were compared for morphology and vascular anatomy of the flower and for embryological development to the stage of mature ovules and pollen. Flowers are liliaceous, the staminate with rudimentary pistils and the pistillate with collapsed anthers. The uncomplicated vascular pattern differs between staminate and pistillate flowers only in the size and degree of maturation of bundles to stamens and carpels. Longer styles appear to be correlated with a greater extent of ovule development in ovaries of staminate flowers. Microsporogenesis in males is normal with wall development corresponding to the Monocotyledonous type. The tapetum is glandular and binucleate, cytokinesis successive, the tetrads isobilateral or occasionally decussate, and the mature pollen grain two-celled. A pair of heteromorphic, possibly sex, chromosomes was observed in heterogametic male plants. Anther development is initially the same in pistillate flowers, but the tapetum degenerates precociously followed by collapse of microspore mother cells. In pistillate flowers the ovules are hemitropous, bitegmic, and slightly crassinucellate. Megasporogenesis-megagametogenesis conforms to the Polygonum type. In staminate flowers ovule development is like that in pistillate flowers until degeneration starts in nucellar and integumentary cells at the chalazal end. Ovules in both homogametic male genotypes rarely complete meiosis, while in the heterogametic males it is normally completed with about one ovule in 20 flowers forming a mature megagametophyte. Since manipulation of sex expression in Asparagus could be important in developing inbred male and female lines for breeding purposes, those aspects of the morphological and embryological observations presented which might be useful in planning experiments to induce sex changes are discussed briefly.     

Morphological variation in the flowers of Jacaratia mexicana A. DC. (Caricaceae), a subdioecious tree

The Caricaceae is a small family of tropical trees and herbs in which most species are dioecious. In the present study, we extend our previous work on dioecy in the Caricaceae, characterising the morphological variation in sexual expression in flowers of the dioecious tree Jacaratia mexicana . We found that, in J. mexicana , female plants produce only pistillate flowers, while male plants are sexually variable and can bear three different types of flowers: staminate, pistillate and perfect. To characterise the distinct types of flowers, we measured 26 morphological variables. Our results indicate that: (i) pistillate flowers from male trees carry healthy-looking ovules and are morphologically similar, although smaller than, pistillate flowers on female plants; (ii) staminate flowers have a rudimentary, non-functional pistil and are the only flowers capable of producing nectar; and (iii) perfect flowers produce healthy-looking ovules and pollen, but have smaller ovaries than pistillate flowers and fewer anthers than staminate flowers, and do not produce nectar. The restriction of sexual variation to male trees is consistent with the evolutionary path of dioecy from hermaphrodite ancestors through the initial invasion of male-sterile plants and a subsequent gradual reduction in female fertility in cosexual individuals (gynodioecy pathway), but further work is needed to confirm this hypothesis.     

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。     

Thrips pollination of the dioecious ant plant Macaranga hullettii (Euphorbiaceae) in Southeast Asia

Discussion about thrips (Thysanoptera) as main pollinators has been controversial in the past because thrips do not fit the preconception of an effective pollinator. In this study, we present evidence for thrips pollination in the dioecious pioneer tree genus Macaranga (Euphorbiaceae). Macaranga hullettii is pollinated predominantly by one thrips species, Neoheegeria sp. (Phlaeothripidae, Thysanoptera). As a reward for pollinators, the protective floral bracteoles function as breeding sites for thrips and trichomal nectaries on the adaxial surface of the floral bracteoles provide alimentation. Flowering phenology of both staminate and pistillate trees was highly synchronized within 3-4 wk periods. In contrast to pistillate trees, staminate trees start to breed the thrips inside the developing inflorescences ～2 wk before anthesis. Breeding of Neoheegeria sp. in the laboratory indicates that the thrips development is completed within ～17 d. Thus, staminate trees offer breeding sites for one thrips generation until the onset of pollen presentation. Intraspecific pollen transfer by thrips was proved by pollen loads of thrips taken from receptive pistillate inflorescences of M. hullettii. Bagging experiments of different mesh sizes showed that seed set reached almost the level of open-pollinated flowers when exclusively tiny insects like thrips were able to enter the net bags, but no apomictic seed set occurred when no insect access was given to the flowers.     

Garcinia brasiliensis: insights into reproductive phenology and sexual system in a Neotropical environment

Sexual systems play a key role in the ecology and evolution of plant populations. Research on Garcinia L. has shown the occurrence of variations in the sexual systems of this genus, with some authors considering it as dioecious, and for some species as gynodioecious or androdioecious. The population of G. brasiliensis in the restinga of Maricá is dioecious with two individuals characterized as monoecious. Anatomical analysis showed that one has pistillate and sterile staminate flowers, being an unusual female and the other has staminate and perfect flowers (functional pollen grains and ovules), being andromonoecious. In addition, field observations, controlled hand-pollinations, reproductive phenology and anatomical analysis showed no evidence of apomixis in this Neotropical species, different from that provided in the literature for paleotropical species. However, asexual reproduction occurs through ramets suggesting that in this environment of restinga the seedlings are subjected to considerable limitations, and vegetative propagation becomes an important alternative for the occupation of the environment. Flowering episodes occurred throughout the study period, featuring a subannual and long-lasting flowering pattern mainly due to multiple blooms of male plants. The floral characters observed in G. brasiliensis are often associated with a generalist pollination system. Apis mellifera and Trigona spinipes are the effective pollinators of G. brasiliensis flowers. Our findings provide new insights into the reproductive biology of Garcinia.     

INTERSPECIFIC GENE FLOW IN CUCURBITA: C. TEXANA VS. C. PEPO

The potential for interspecific genetic exchange was examined by monitoring flowering patterns, pollinator movement, and gene flow among experimental populations of the Texas gourd (Cucurbita texana) and cultivars of Cucurbita pepo. While flowering patterns and pollinator movement tended to maximize self-pollination and local gene exchange, movement of effective pollen exceeded 1,300 m. This movement, mediated by the solitary bee Xenoglossa strenua and monitored by tracking allozyme variants, produced interspecific hybrids in 5% of the progeny from experimental plants. Interspecific gene exchange occurred in either direction with either species serving as staminate or pistillate parent. No obvious constraints to gene flow among plants representing C. texana and distinctive cultivars (vars. ovifera, medullosa, melopepo) of C. pepo were detected. Genetic exchange among different species and cultivars is enhanced by the foraging behavior of Xenoglossa. Multiple visits to either staminate (pollen carryover) or pistillate (multiple pollinations) flowers often result in the deposition of mixed pollen on receptive stigmas. The wild type (C. texana) can donate and receive effective pollen when growing under both weedy and natural conditions. The observed lack of interspecific reproductive isolation supports treatment of cultivars and wild types as a single species and, in conjunction with available data concerning temporal/geographical relationships among bees, squash, gourds, and humans in eastern North America, suggests the possibility of long-term genetic interaction between wild types and domesticates.     

Attalea phalerata and Bactris glaucescens (Arecaceae, Arecoideae): Phenology and pollination ecology in the Pantanal, Brazil

We compared the reproductive and vegetative phenology and pollination ecology of sympatric Attalea phalerata and Bactris glaucescens palms (Arecaceae) in the Pantanal, Brazil, in a riparian forest area subject to periodic flooding. Attalea phalerata has a solitary stem and produces staminate, pistillate and rarely bisexuals inflorescences that open during the day. Bactris glaucescens has multiple stems and has bisexual flowers with crepuscular/nocturnal anthesis. Both species present bud break and leaf-budding during the year. Attalea phalerata shows continual annual flowering with ripening of fruits during the dry season. For B. glaucescens flowering occurs simultaneously with fruiting for about seven months, and fruit production can be influenced by temperature and level of flooding. The difference in the timing of anthesis of the flowers ensures production of floral resources during both day and night when both species of palms are flowering. The floral structures of both species have morphological characteristics associated with pollination by insects (mainly beetles), such as the presence of odor, stigmatic secretion, heat production, and adhesive substances (“pollenkitt”) in the pollen. In A. phalerata, the main pollinators were Mystrops sp. (Nitidulidae) and Madarini (Curculionidae). Derelomus sp. (Curculionidae) and Paratenetus sp. (Tenebrionidae) beetles visited B. glaucescens during the day and could have pollinated the flowers at these occasions.     

Growth, Sex Expression and Yield of Squash Melon (Citrullus vulgarisvar. fistulosus) as Influenced by 2(chloroethyl)phosphonic acid, 2,3,5-tri-iodobenzoic acid and (2-chloroethyl) trimethyl-ammonium chloride

Single aqueous sprays of 2(chloroethyl)phosphonic acid (CEPA) 250, 500 and 1000 mg/1; 2,3,5-triiodobenzoic acid (TIBA) 25, 50 and 100 mg/1; and (2-chloroethyl) trimethyl-ammonium chloride (CCC) 250, 500 and 1000 mg/1 were applied to squash melon (Citrullus vulgaris Schrad. var. fistu-losus Stocks.) at the 2–3 leaf stage. Though the final length of the main axis and number of lateral branches were not affected by any treatment, CEPA retarded growth of young plants by reducing the internodal length. It did not change the flowering pattern but delayed flowering and reduced the production of both pistillate and staminate flowers. However, it increased the per cent femaleness as a result of comparatively more suppression of staminate flowers. The TIBA 25 and 50 mg/1 and CCC 500 mg/1 did not affect the staminate flower production but increased the pistillate flowers, which increased the per cent femaleness. The CEPA decreased while both TIBA 25 and 50 mg/1, and CCC 500 mg/1 increased the number of fruits per plant and the yield. The mode of action of the chemicals has been discussed.     

Reproductive structures of an extinct platanoid from the early Cretaceous (latest Albian) of eastern North America

Two new species of platanoid reproductive structure are described from the Bull Mountain locality in the Patapsco Formation (Potomac Group) of northeastern Maryland, USA. Pistillate inflorescences and infructescences (Platanocarpus elkneckensis sp. nov.) consist of flowers and fruits in sessile globose heads that are borne on an elongate axis. Individual pistillate flowers consist of five free carpels surrounded by prominent tepals. Staminate inflorescences, flowers and isolated stamens are assigned to Hamatia elkneckensis gen. et sp. nov. Staminate flowers are borne in a globose head with a small number of stamens (five?) per flower. Stamens consist of very short filaments, long anthers with strongly valvate dehiscence and an apically expanded connective. The connective expansion is frequently very well-developed, hook-like and extends down the ventral surface of the stamen. Anthers contain small, tricolporate, reticulate pollen. Association evidence, similarity of inflorescence structure and the occurrence of Hamatia-type pollen on flowers, carpels and fruits of Platanocarpus elkneckensis suggests that the staminate and pistillate material was produced by a single species of plant. The “Hamatia-plant” provides further evidence of pentamerous floral structure in mid-Cretaceous platanoids and documents the occurrence of unequivocal tricolporate pollen in the platanoid complex.     

HETERODICHOGAMY IN GRAYIA BRANDEGEI (CHENOPODIACEAE): REPORT FROM A NEW FAMILY

The reproductive biology of Grayia brandegei was examined. Although monoecious, Grayia brandegei exhibits a phenotypic dimorphism of protogynous and protandrous mating types known as heterodichogamy. For individual plants, the temporal separation of staminate and pistillate flowering phases appears to be complete. No self-fertilization is possible in protandrous plants, but may be possible in protogynous plants provided pistillate flowers remain unfertilized. Flowering phases of protogynous and protandrous mating types are synchronized and reciprocal, ensuring cross-fertilization between mating types. Less than 15% overlap of sexual functions occurred between plants of the same mating type. Protogynous and protandrous plants were randomly dispersed in the environment and in relation to each other. Mating type frequencies did not differ significantly from 1:1. We discuss the possibility of a heterodichogamous pathway to dioecy.