STRUCTURE AND DEVELOPMENT OF THE PERIANTH IN DOWNINGIA BACIGALUPII

The structure and ontogeny of the calyx and corolla of Downingia bacigalupii Weiler (Campanulaceae; Lobelioideae) were investigated for the purpose of comparing perianth development with previous observations on the floral bract, as well as elucidating the mechanism of development of the zygomorphic, sympetalous corolla. Sepals are uni-traced with a palmate, reticulate venation. They have basal and apical hydathodes, as well as storage tracheids. Sepals show a reduction in size, venation and hydathode number when compared to the bract. The pentamerous, zygomorphic corolla is bilabiate, consisting of a three-lobed adaxial lip and a two-lobed abaxial lip connected by a short tubular region. The constituent petal lobes are also uni-traced and have a reticulate venation, resembling that of the sepal and bract, but lack storage tracheids and hydathodes. Sepals arise in an adaxial to abaxial succession and are initiated in the outer corpus layer of the floral apex. Expansion of the floral apex follows and is accompanied by the establishment of a second tunica layer. Sepals undergo apical, marginal, and intercalary growth accompanied by acropetal differentiation of procambium. The petals arise simultaneously and are initiated in the second tunica layer and the outer corpus cells. After initiation, the petals exhibit a period of apical and marginal growth followed by intercalary growth. Apical growth in petals is less protracted than in sepals, but plate meristem activity is more extensive. The free petal lobes become temporarily fused by an interlocking of marginal epidermal layers, but they separate at anthesis. Zonal growth beneath the originally free lobes forms the tube and lip regions of the sympetalous corolla. Zygomorphy is evident from the time of initiation of petals and is accentuated by later differential growth. Comparative observations of corolla ontogeny in autogamous species of Doumingia indicate that the reduced corollas in these taxa are derived by a simple process of neoteny.     

南瓜两性花的形态与结构研究

对南瓜的两性花进行外部形态及结构的研究,结果表明：南瓜的两性花可有子房上位花、子房半下位花和子房下位花三种类型：花萼、花冠均为５,合瓣;雄蕊３,其中有两枚各由两个雄蕊合生而成,分离和部分联合,花药结构特殊,花粉发育正常;雌蕊具有单个或两个柱头及花柱。子房壁结构正常。上方的子房壁表皮发育完好,具气孔器。胚珠在外形上也发育正常。两性花与单性花同生长在一个植株上,可以连续多代稳定地遗传,萌生当代植株的种子来源于上一代雌花。这对探讨被子植物的系统进化关系有着重要的意义     

Floral morphology of the family compositae

Vernonia anthelmintica (Linn.) Willd., tribe Vernonieae, family Compositae has been studied. The cypsela has dimorphic pappus, 20 longitudinal ribs (10 primary and 10 secondary), and two types of hairs developing from single epidermal cells on surface. Certain features of vasculature of floret observed in this taxon have been considered as intermediate between relatively primitive and highly advanced conditions known in the family. Others elucidate further that the ovary is bilocular with axile placentation at base in which region the ovule is attached, and unilocular with parietal placentation in the remaining portion, and also that the ovule possibly belongs to one of the two carpels, which alone is fertile, and is an organ morphologically double in nature. The pappus has been regarded as a longitudinally dismembered calyx tube. The tissue of the ovule outside nucellus differentiates into an endothelium, a periendothelial zone, and an outer zone. In the mature seed the thickned outer epidermis and remmants of a few subjacent layers form the outer zone, and endothelium redued to a pellicle, the inner zone of the seed coat. The jacket layer of the endosperm persists between the seed coat and the dicotyledonous, orthorrhizal embryo. The pericarp development resembles that of most other Compositae in an early differentiation of the ovary wall into two zones, and later breakdown of a major part of the inner zone. But it is quite characteristic in other respects, especially, the peripheral chin of firovascular bundles which forms its main mechanical zone.     

MORPHOLOGY AND ANATOMY OF FLORAL AND EXTRAFLORAL NECTARIES IN CAMPSIS (BIGNONIACEAE)

The genus Campsis (Bignoniaceae), with one New World and one Old World species, is unusual among temperate plants in having five distinct nectary sites. Multiple nectaries occur at all four of the extrafloral sites (petiole, calyx, corolla, fruit), representing an advanced strategy for ant attraction. The morphology and anatomy of the extrafloral nectaries in both species are uniform for the petioles, calyces, and young fruits; those on the outer corolla lobes are of slightly different forms. The generalized structure consists of one layer of basal cells, and a one- to two-layered secretory cup. Because of their small size, there is no vascular tissue in them. The large, vascularized (phloem only) floral nectary is an annular structure subtending the ovary.     

弯齿盾果草(紫草科)花序及花的形态发生

以弯齿盾果草不同发育时期的花芽为材料,在体视显微镜解剖观察的基础上使用扫描电镜对弯齿盾果草花序、花及果实的发育过程进行了观察。结果显示:(1)弯齿盾果草的花序是由最初的一个球形花序原基经过多次分裂形成的,且花序发生式样符合蝎尾状聚伞花序结构,而非通常所描述的镰状或螺状聚伞花序;花序发生过程中无单一主轴,花序轴是由侧枝连接而成,每一朵花原基有其对应的1枚苞片,下一花原基是从相邻的上一枚苞腋里发生,相邻两花原基交错互生。(2)花器官的发生是按照花萼原基、花冠原基、雄蕊原基和雌蕊原基的顺序发育,但雄蕊原基的花药部分发育速度要比花冠原基快,所以花器官的发育是按照花萼、雄蕊、花冠和雌蕊的顺序发育。(3)子房四深裂结构是由4个原基分别发育,而后相互靠拢而成。(4)小坚果表面的附属结构发生于子房发育后期,其背面的内外层突起分别是由生长较快的外部组织的边缘通过上部内缩和下部向外环状生长形成。     

Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb

The annual Mediterranean herbTheligonum cynocrambe shows a peculiar combination of morphological characters, e.g., switch from decussate to spiral phyllotaxis with 90–100° divergence, combined with a change from interpetiolar to lateral stipules, anemophily, lack of calyx, flowers often dimerous to trimerous, corolla fused in both male and female flowers, male flowers extra-axillary, with 2–19 stamens per flower, female flowers axillary, with inferior uniovulate ovary, basilateral style and perianth, nut-like fruits with elaiosome. In male flowers the androecium emerges as an (uneven) elliptical rim with a central depression. This common girdling primordium is divided up into several stamen primordia. In male flowers with low stamen number the stamen primordia may occupy the corners alternating with the corolla lobes. There are no epipetalous androecial primordia that secondarily divide into stamens. Male flowers occasionally show a hemispherical base that may be interpreted as remnant of the inferior ovary. In female flowers a ring primordium grows into a tube on which the petal lobes arise. The perianth and style become displaced adaxially by uneven growth of the inferior ovary. The ovary is basically bilocular. The lower region of the ovary is provided with a septum that is overtopped and hidden by the single curved ovule.Theligonum is referred to theRubiaceae-Rubioideae, with theAnthospermeae andPaederieae as most closely related tribes.     

甘蔗雌雄蕊的解剖学研究

甘蔗雄蕊花药的壁由四层细胞构成,药室内壁在花药成熟时才发育。花粉母细胞减数分裂属连续型,四分体正常发育为小孢子,但当开花时,只有少数进一步发育为成熟的花粉粒。甘蔗雌蕊柱头是分枝的,每一柱头毛是由数个细胞质浓密的细胞,呈不规则的纵列而组成。倒生胚珠一个,着生于子房室的内侧;内、外珠被均由两层细胞构成,外珠被上侧未能生长到珠孔,内珠被的内层细胞在胚珠发育中不断增大体积。石蜡制片是以樟油为透明剂的。     

Ovary and Gametophyte Development Are Coordinately Regulated by Auxin and Ethylene following Pollination

The differentiation and development of ovules in orchid flowers are pollination dependent. To define the developmental signals and timing of critical events associated with ovule differentiation, we have examined factors that regulate the initial events in megasporogenesis and female gametophyte development and characterized its progression toward maturity and fertilization. Two days after pollination, ovary wall epidermal cells begin to elongate and form hair cells; this is the earliest visible morphological change, and it occurs at least 3 days prior to pollen germination, indicating that signals associated with pollination itself trigger these early events. The effects of inhibitors of ethylene biosynthesis on early morphological changes indicated that ethylene, in the presence of auxin, is required to initiate ovary development and, indirectly, subsequent ovule differentiation. Surprisingly, pollen germination and growth were also strongly inhibited by inhibitors of ethylene biosynthesis, indicating that male gametophyte development is also regulated by ethylene. Detailed characterization of the development of both the female and male gametophyte in pollinated orchid flowers indicated that pollen tubes entered the ovary and grew along the ovary wall for 10 to 35 days, at which time growth was arrested. Approximately 40 days after pollination, coincident with ovule differentiation as indicated by the presence of a single archesporial cell, the direction of pollen tube growth became redirected toward the ovule, suggesting a chemical signaling between the developing ovule and male gametophyte. Taken together, these results indicate that both auxin and ethylene contribute to the regulation of both ovary and ovule development and to the coordination of development of male and female gametophytes.     

中国夹竹桃目的形态演化趋势及其进化系统

本文探讨了中国夹竹桃目主要器官分类性状的进化趋势,发现了中国夹竹桃目有关花萼、花冠、副花冠、雄蕊、花粉、子房和果实等分类性状密切关系和演化的规律性(图1),从而在作者参考蒋英教授工作成就的基础上,提出了中国夹竹桃目各分类群的进化系统(图2)。     

Patterns and Significance of Floral Development in Whytockia (Gesneriaceae)

Abstract: The floral development of Whytockia W. W. Smith has been studied in order to explore the developmental basis for the arrangement and differentiation patterns of floral organs, and the evolutionary relationship between Whytockia and allies in floral development. The descending imbricate aestivations in both calyx and corolla have remarkably different ontogenetic patterns between calyx and corolla which are derivative with respect to the development of the valvate aestivations in the four-stamened Rhynchoglossum. Both corolla lobes and stamens are initiated simultaneously from the same ring meristem. However, the five stamens remarkably precede the initiation of the five corolla lobes. Also, the adaxial stamen is suppressed after initiation to become a staminode, concomitant with retardation of its adjacent organs during development. This situation, together with the non-acropetal order among whorls of floral organs in Whytockia, is possibly related to a late expression and a remarkably different expression pattern of cycloidea- like genes as compared to Antirrhinum. Furthermore, the axile placentation in the bilocular ovary of Whytockia is formed by an involute closure of carpels rather than derived from a secondary fusion of two intrusive parietal placentae.     

大叶补血草的大、小孢子发生与雌、雄配子体的发育

系统地报道了大叶补血草(Limonium gmelinii (Willd.) Kuntze)的大、小孢子发生和雌、雄配子体的形成发育过程。主要结果如下：(1)小孢子母细胞减数分裂过程中的胞质分裂为同时型,四分孢子多为正四面体形, 也有少数为左右对称形;(2)成熟花粉为三细胞型,具3个萌发孔;(3)花药壁由5层细胞组成,最外层为表皮,其内分别为药室内壁、中层、绒毡层,绒毡层为变形型,花药壁的发育属于基本型;(4)大叶补血草的雌蕊由5心皮合生,子房1室,基生胎座,胚珠1个,拳卷型,双珠被,厚珠心;(5)孢原细胞发生于珠心表皮下,经一次平周分裂,形成造孢细胞,由造孢细胞直接发育成大孢子母细胞,大孢子母细胞减数分裂形成4个大孢子呈直线排列,合点端大孢子具功能,属于典型的蓼型胚囊发育。     

巨龙竹生殖器官形态结构及雌、雄配子体的发育

通过石蜡切片的方法对巨龙竹生殖器官结构、大小孢子的发生和雌、雄配子体的发育过程进行了观察研究。 巨龙竹为一心皮组成的单室单子房,子房内具有一个胚珠,倒生、双珠被、厚珠心。大孢子母细胞减数分裂形成线形排列的4个大孢子,合点端大孢子具功能。胚囊的发育为蓼型,具多个反足细胞。巨龙竹的花药壁由4层结构组成,包括表皮、药室内壁、中层、绒毡层;花药壁发育为单子叶型,绒毡层为腺质型。小孢子母细胞减数分裂中的胞质分裂为连续型,四分孢子为四面体型;成熟花粉粒为2细胞型,具1个萌发孔。小穗发育雌雄异熟,雌蕊的发育早于雄蕊的发育。     

柽柳大、小孢子发生和雌、雄配子体发育的观察

利用常规石蜡制片技术,对柽柳(Tamarix chinensis Lour.)的大、小孢子发生及雌、雄配子体发育过程进行了观察。主要结果如下:(1)花药壁由五层细胞组成,从外向内分别为表皮、药室内壁,两层中层和绒毡层。药壁的发育属于基本型。绒毡层为分泌型。(2)孢原细胞为多孢原起源。小孢子母细胞减数分裂过程中的胞质分裂为连续型,形成的四分孢子为四面体型;同一药室的小孢子母细胞减数分裂几乎完全同步。(3)成熟花粉粒为2细胞型,具3个萌发孔。(4)柽柳为三心皮构成的单室复子房,每子房具有10~20个胚珠,基底胎座,胚珠为双珠被、厚珠心、倒生型。大孢子母细胞减数分裂形成1+3排列的4个大孢子, 4个大孢子全部参与胚囊的形成。(5)胚囊发育为贝母型,反足细胞在胚囊成熟时充分发育。(6)同一朵花中,前期雄蕊的发育早于雌蕊的发育,后期当花粉成熟时,雌配子体也达到成熟,雌雄蕊发育趋于同步。     

DEVELOPMENT OF THE SEED OF SOLANUM PHUREJA

Dnyansagab , Vishnu R., and Delmer C. Cooper . (U. Wisconsin, Madison.) Development of the seed of Solanum phureja. Amer. Jour. Bot. 47(3) : 176—186. Illus. 1960.—Ontogeny of the seed of Solanum phureja Juz. et Buk. is described. The megagametophyte, during the course of its development, ruptures the nucellus and at maturity lies in direct contact with the inner layer (endothelium) of the single massive integument. The mature megagametophyte, a 7-celled structure, consists of a 3-celled egg apparatus, an endosperm mother cell with fused polar nuclei and 3 persistent antipodals. Both 2- and 3-celled mature pollen grains are formed within anthers of the same flower; hence this character cannot be considered of any taxonomic value. Double fertilization occurs between 24 and 72 hr. after pollination. A cellular endosperm is formed, the peripheral layer acting as an absorbing tissue during the early ontogeny of the seed. Later this layer becomes organized as an aleurone layer and thereafter the source of nutrients is via the basal portion of the endosperm immediately adjacent to the apical end of the vascular tissue of the developing seed. Embryo development follows the Nicotiana variation of the Solanad type. The mature testa is composed of an outer layer of thick-walled epidermal cells, an inner layer of thin-walled cells and an intervening mass of disorganized tissue. In those instances where the ovule or young seed aborts, the endothelial cells of the integument become hyperactive and proliferate to such an extent that the space formerly occupied by the gametophyte or the developing endosperm and embryo becomes completely filled with endothelial tissue.     

HISTOGENESIS OF THE ANDROECIUM AND GYNOECIUM IN DOWNINGIA BACIGALUPII

A histogenetic investigation of the synandrous androecium and syncarpous gynoecium in the flower of Downingia bacigalupii Weiler (Campanulaceae; Lobelioideae) was undertaken for the purpose of comparing the modes of initiation, early growth and fusion in these floral whorls with that reported previously for the perianth in this species. Stamens are initiated as separate organs from the second tunica layer and underlying corpus regions of the concave floral meristem. Subsequent growth of stamens involves apical and intercalary growth in length and rudimentary marginal growth in breadth. Tissues of the four microsporangia originate from hypodermal sporangial initial cells and the filament is formed by intercalary growth at the base of the anther. Lateral fusion of stamens is ontogenetic and involves cuticular fusion of adjacent epidermal layers. The two emergent carpel primordia arise as crescentic organs by periclinal divisions in the second tunica layer and corpus zones. Carpel primordia also undergo apical and intercalary growth in length as well as extensive marginal growth in breadth. Radial growth in carpels is mediated by an adaxial meristem which shows its greatest concentration of activity at the carpel margins. Carpel fusion appears to be partially ontogenetic accompanied by zonal growth. Closure of the stylar canal is by the formation of a transmitting tissue derived from the protodermal layers of the adaxial carpel surfaces. A discoid nectary is initiated around the base of the style and formation of the inferior ovary is by intercalary growth of the base of the concave floral bud. The two parietal placentae originate as longitudinal outgrowths from the walls of the floral cup. Ovule initiation is simultaneous at first and then intercalary during subsequent elongation of the ovary. The ovules are anatropous, unitegmic and tenuinucellate. Stamen and carpel procambium shows a slight delay in differentiation when compared to that reported for the perianth and bract, but in all other respects carpels resemble other floral organs in their patterns of histogenesis and early growth. Stamens diverge from the other floral organs in their early pattern of growth, but a consideration of all features of their histogenesis suggests an appendicular rather than an axial interpretation of these organs.     

藏药椭圆叶花锚的胚胎学

本文首次报道了藏药椭圆叶花锚的胚胎学特征。花药四室,药壁发育为双子叶型;绒毡层异型起源,腺质型绒毡层。小孢子母细胞减数分裂为同时型,四分体的排列方式为四面体型;成熟花粉为３－细胞。子房２心皮,而二心皮连接处强烈膨大、内凸,４列胚珠。薄珠心,单珠被,直生胚珠。胚囊发育为蓼型。胚乳发育为核型。胚胎发育为茄型酸浆Ｉ变型。反足细胞在胚囊成熟时期宿存。承珠盘发达。果实成熟时,种子只发育至球型胚阶段。比较了该种与龙胆族其它属、种的胚胎学特征,发现它们大部分特征是相似的,但在如下３个特征上存在区别：子房二心皮连接处强烈膨大、内凸;直生胚珠;具有发达的承珠盘。其胚胎学特征的系统学和分类学意义有待进一步比较与评价     

POSTGENITAL CARPEL FUSION IN CATHARANTHUS ROSEUS (APOCYNACEAE) IV. SIGNIFICANCE OF THE FUSION

The ontogenetic and phylogenetic significances of the postgenital carpel fusion in Catharanthus roseus were investigated experimentally. When one of the two carpel primordia was surgically destroyed prior to carpel fusion, the remaining carpel differentiated nevertheless; however, the tissue development on the adaxial (normally fused) carpel face appeared aberrant. Possible reasons for this modified ontogeny are discussed. The suggestion that syncarpy is phylogenetically significant because any pollen tube can potentially reach any ovule was verified in C. roseus by monitoring pollen tube growth following spatially controlled pollinations. The suture formed by the postgenital fusion was no barrier to pollen tube growth, and the pollen tubes rapidly became distributed throughout the stigmatic tissue.     

DYNAMICS OF POLLEN TUBE GROWTH IN THE WILD RADISH RAPHANUS RAPHANISTRUM (BRASSICACEAE). II. MORPHOLOGY,CYTOCHEMISTRY AND ULTRASTRUCTURE OF TRANSMITTING TISSUES,AND PATH OF POLLEN TUBE GROWTH

The relative importance of prezygotic mechanisms of gametophytic competition and selection are often unclear due to an absence of observations on the gynoecium and pollen tube growth in vivo. We used LM, SEM, and TEM to study the structure of the gynoecium and the path of pollen tube growth in Raphanus raphanistrum, a sporophytically self-incompatible annual. Wild radish has a papillate stigma and a solid style. A septum, which is characteristic of cruciferous gynoecia, is present in the ovary. After germination on the stigma, pollen tubes grow in the secretion of the transmitting tract of the style. The stylar secretion stains positive for acidic polysaccharides and insoluble carbohydrates, and negative for lipids and protein. In the ovary, the transmitting tissue is contained inside the septum. The secretion in the ovary stains positive only for acidic polysaccharides. Pollen tubes travel inside the septum as they enter the ovary and must exit to the surface of this tissue before ovule fertilization can occur. Pollen tube growth on the septum tracks the intercellular junctions of the septum epidermis where the secretion leaks out through a torn cuticle. Tubes must grow across the obturator before reaching the micropyle of an ovule. The temporal pattern with which tubes growing into the ovary exit the septum can contribute to the previously observed nonrandom patterns of fertilization (Hill and Lord, 1986).