蓝堇草属(毛茛科)花形态发生的扫描电子显微镜观察

花的发生和发育过程研究可以发现早期进化的轨迹,为系统发育的研究提供重要线索。蓝堇草属(Leptopyrum)为毛茛科唐松草亚科一单种属,仅包含蓝堇草一种,其花的发生和发育过程仍为空白。为了深入理解唐松草亚科乃至毛茛科花发育多样性和演化规律,该文运用扫描电子显微镜(SEM)观察了蓝堇草各轮花器官的形态发生和发育过程。结果表明:该属植物所有的萼片、花瓣、雄蕊和雌蕊均为螺旋状发生,花器官排列式样也为螺旋状; 5枚萼片原基宽阔,5枚花瓣原基圆球形、位于萼片原基的间隔,且在后期表现为延迟发育现象,雄蕊原基较小、为圆球形; 花瓣原基和雄蕊原基连续发生,无明显的时空间隔,但与萼片原基有时空间隔; 心皮原基为马蹄形对折,柱头组织由单细胞乳突组成; 胚珠倒生、具单珠被。该属花器官螺旋状排列、胚珠具单珠被在唐松草亚科中是独有的性状,花发育形态学证据支持了该属的特殊性。     

One size fits all? Molecular evidence for a commonly inherited petal identity program in Ranunculales

Petaloid organs are a major component of the floral diversity observed across nearly all major clades of angiosperms. The variable morphology and development of these organs has led to the hypothesis that they are not homologous but, rather, have evolved multiple times. A particularly notable example of petal diversity, and potential homoplasy, is found within the order Ranunculales, exemplified by families such as Ranunculaceae, Berberidaceae, and Papaveraceae. To investigate the molecular basis of petal identity in Ranunculales, we used a combination of molecular phylogenetics and gene expression analysis to characterize APETALA3 (AP3) and PISTILLATA (PI) homologs from a total of 13 representative genera of the order. One of the most striking results of this study is that expression of orthologs of a single AP3 lineage is consistently petal-specific across both Ranunculaceae and Berberidaceae. We conclude from this finding that these supposedly homoplastic petals in fact share a developmental genetic program that appears to have been present in the common ancestor of the two families. We discuss the implications of this type of molecular data for long-held typological definitions of petals and, more broadly, the evolution of petaloid organs across the angiosperms.     

The floral development and floral anatomy ofChrysosplenium alternifolium, an unusal member of the Saxifragaceae

The floral development and anatomy ofChrysosplenium alternifolium were studied with the scanning electron microscope and light microscope to understand the initiation sequence of the floral organs and the morphology of the flower, and to find suitable floral characters to interpret the systematic position of the genus within the Saxifragaceae. The tetramerous flower shows a highly variable initiation sequence. The median sepals and first stamens arise in a paired sequence resembling a dimerous arrangement, but the first sepal and stamen arise on the side opposite to the bract. Transversal sepals and stamens emerge sequentially, as one side often precedes the other; sepals and stamens occasionally arise on common primordia. Initiation of the gynoecium is more constant with two median carpel primordia arising on a sunken floral apex. Several flowers were found to be pentamerous with a 2/5 initiation sequence. Flowers were invariably found to be apetalous without traces of petals in primordial stages; this condition is interpreted as an apomorphy. It is postulated that the development of a broad gynoecial nectary is responsible for the occurrence of an obdiplostemonous androecium. The gynoecium shows a number of anatomical particularities not observed in other Saxifragaceae. The presence and distribution of colleters is discussed.     

Floral Morphology and Relationships of Clusia gundlachii with a Discussion of Floral Organ Identity and Diversity in the Genus Clusia

The genus Clusia L. is highly variable in many floral features. Several Clusia species have floral organs of mixed or uncertain identity, such as organs that are transitional between bracteoles and sepals, petaloid sepals, and partly petaloid stamen rings. Unique in Clusia is the "corona" of Clusia gundlachii Stahl, a thick, urn-shaped structure that is initiated as a ring primordium. In male flowers it surrounds a synandrium, and in female flowers it surrounds the ovary and a row of staminodes. The corona combines features typical of both petals and stamens of other Clusia species. It is hypothesized that this corona may be the result of the altered expression patterns of the genes that determine floral organ identity. Clusia gundlachii has many floral features in common with two small genera that are sometimes included in Clusia: Havetiopsis and Oedematopus. These genera have four thick connivent petals. Their apparent close relationship makes it seem likely that the corona of C. gundlachii evolved via congenital fusion of such petals. The corona is also somewhat similar to the staminodial rings present in many Clusia species, but taxa in which such organs occur show little similarity to C. gundlachii in terms of other floral characters.     

Effect of lateral suppressor on petal initiation in tomato

Flowers developing on tomato ( Lycopersicon esculentum ) plants homozygous for the lateral suppressor ( ls ) mutation lack petals. Scanning electron micrographs revealed that in ls plants no second whorl organs were initiated. The initiation of first, third, and fourth whorl organs were unaffected by this mutation. To investigate interactions between the cells in different layers of the floral meristem during organ initiation, a periclinal chimera between wild-type and ls tomato was generated. Flowers of the chimera having ls cells in the outer meristem layer (L1) and wild-type cells in internal layers (L2 and L3) developed normally, including the initiation of organ primordia that differentiated as petals in normal positions within the second whorl. L1 of the chimera developed in a non-autonomous manner during petal development. Thus, wild-type cells occupying the internal meristem layers provided developmental cues necessary for initiation of petal primordia at appropriate positions on the floral meristem. L1 cells carrying the lateral suppressor mutation were fully capable of responding to this information and differentiated appropriately.     

A SCANNING ELECTRON MICROSCOPIC STUDY ON THE INITIATION AND DEVELOPMENT OF FLORAL ORGANS OF BRASSICA NAPUS (CV. WESTAR)

The initiation and development of the floral organs of Brassica napus L. (cv. Westar) were examined using the scanning electron microscope. After transition of the vegetative apex into an inflorescence apex, flower primordia were initiated in a helical phyllotactic pattern. The sequence of initiation of the floral organs in a flower bud was that of sepals, stamens, petals and gynoecium. Of the four sepal primordia, the abaxial was initiated first, followed by the two lateral and finally the adaxial primordium. The four long stamens were initiated simultaneously in positions alternating with the sepals. The two short stamens were initiated basipetal to and outside the long stamens, and opposite the lateral sepals. The petals arose on either side of the two short stamens and the gynoecium was produced from the remainder of the apex. During development, the sepal primordia curved sharply at the tips and tightly enclosed the other organs. Stamen primordia developed tetralobed anthers at an early stage while filament elongation occurred just prior to anthesis. A unique pattern of bulbous cells was present on the abaxial surface of the anther. Growth of petal primordia lagged relative to the other floral organs but expansion was rapid prior to anthesis. The gynoecium primordium was characterized by an invagination early in development. At maturity, there was differentiation of a papillate stigma, an elongated style and a long ovary marked externally by sutures and divided internally by a septum. Distinct patterns of cuticular thickenings were observed on the abaxial and adaxial surfaces of the petals and stamens and on the surface of the style. The patterns were less obvious on the sepals and ovary. Stomata were present on both surfaces of the mature sepals, on the style and restricted areas on the abaxial surface of the anthers and nectaries but were absent from the petals, the adaxial surface of the stamens and the ovary. No hairs were present on any of the floral organs.     

NORMAL FLORAL ONTOGENY AND COOL TEMPERATURE-INDUCED ABERRANT FLORAL DEVELOPMENT IN GLYCINE MAX (FABACEAE)

Floral onset in soybean (Glycine max cv. Ransom) is characterized by precocious initiation of axillary meristems in the axils of the most recently initiated leaf primordium. During floral transition, leaf morphology changes from trifoliolate leaf with stipules, to a three-lobed bract, to an unlobed bract. Soybean flowers initiated at 26/22 C day/night temperatures are normal, papilionaceous, and pentamerous. Sepal, petal, and stamen whorls are initiated unidirectionally from the abaxial to adaxial side of the floral apex. The median sepal is located abaxially and the median petal adaxially on the meristem. The organogeny of ‘Ransom’ flowers was found to be: sepals, petals, outer stamens plus carpel, inner stamens; or, sepals, petals, carpel, outer stamens, inner stamens. The outer stamen whorl and the carpel show possible overlap in time of initiation. Equalization of organ size occurs only within the stamen whorls. The sepals retain distinction in size, and the petals exhibit an inverse size to age relationship. The keel petals postgenitally fuse along part of their abaxial margins; their bases, however, remain free. Soybean flowers initiated at cool day/night temperatures of 18/14 C exhibited abnormalities and intermediate organs in all whorls. The gynoecium consisted of one to ten carpels (usually three or four), and carpel connation varied. Fusion of keel petals was often lacking, and stamen filaments fused erratically. Multiple carpellate flowers developed into multiple pods that were separate or variously connate. Intermediate type organs had characteristics only of organs in adjacent whorls. These aberrant flowers demonstrate that the floral meristem of soybean is not fixed or limited in its developmental capabilities and that it has the potential to produce alternate morphological patterns.     

Comparative floral ontogeny in Detarieae (Leguminosae: Caesalpinioideae). 2. Zygomorphic taxa with petal and stamen suppression

Marked floral zygomorphy and a reduced number of petals and/or stamens are the character traits that distinguish the taxa described (species of Afzelia, Berlinia, Gilbertiodendron, Macrolobium, Neochevalierodendron, Paramacrolobium, Phyllocarpus, and Tetraberlinia). All have an "Omega"-shaped floral apex after bracteole initiation, bracteoles large when initiated, helical sepal initiation, unidirectional petal initiation (simultaneous in Afzelia, not determinable in Tetraberlinia), and unidirectional stamen initiation. Floral zygomorphy is expressed primarily by one petal being much larger than the others and by suppression of several of the stamens. Five petals are initiated in all; suppression begins in late development. Either two petals (Neochevalierodendron, Phyllocarpus) or four petals (Afzelia, Berlinia, Macrolobium, Tetraberlinia) are suppressed. All ten stamens are initiated; at midstage, suppression begins in either three stamens (Afzelia) or seven stamens (Gilbertiodendron, Macrolobium, Paramacrolobium). Other expressions of zygomorphy may include diadelphy, stamen filament connation late in development, or displacement of the carpel from a central position to the adaxial side of the hypanthium. There is no loss of organs similar to that which occurs in some other Detarieae.     

UNIDIRECTIONAL ORGAN INITIATION IN LEGUMINOUS FLOWERS

The order of initiation of floral organs is compared in several legumes. In Bauhinia fassoglensis, a caesalpinioid, the sepals are initiated helically, with the first one forming abaxially. In Genista tinctoria and Lupinus affinis (both papilionoids) the sepals are initiated unidirectionally, with the first forming on the abaxial side of the floral apex and subsequent sepals initiating laterally and then adaxially. All three taxa show unidirectional order of initiation for petals, first-whorl stamens, and second-whorl stamens. In each whorl, the first member or members form on the abaxial side, next to the subtending bract, then the lateral ones, and lastly the member(s) on the adaxial side, next to the axis. In Lupinus and Genista there are overlaps in time of initiation between organs in different whorls; for instance, the first stamens begin initiating before the last petals appear. Size differences among members of a whorl are evident in early stages, but may disappear after organogeny ceases, when the members become equal in size in each whorl. This precocious onset of dorsiventrality in floral development is viewed as a specialized feature.     

贵州特有药用植物毕节小檗的形态变异

为探讨贵州特有药用植物毕节小檗(Berberis guizhouensis)形态多样性的变异式样及其适应机制,对其野外居群营养器官和花部形态进行研究。对毕节小檗一直空缺的花部性状进行了描述。结果表明,毕节小檗的性状在同一居群或同一性状在不同居群中都存在较丰富的变异,变异系数为0.03~0.57。从总体来看,营养器官性状的变异程度要高于花部性状。单因素方差分析表明居群间部分花部性状存在显著差异(P0.05)。聚类分析表明小白岩居群形成1个独立的分支,这种分化很可能是环境压力选择的结果。主成分分析表明花部性状与营养性状在维持形态结构的稳定性具有协调一致的生态适应。     

Synthèse et hydrolyse du saccharose dans l'œillet alimenté avec une solution glucosée

Sucrose synthesis and sucrose hydrolysis in cut carnations (Dianthus caryophyllus) supplied with glucose solution .
Sucrose synthesis and hydrolysis in cut carnations supplied with a glucose solution have been investigated using both entire floral branches and floral branches without any of their organs (petals, non-petaloid pieces, leaves) and isolated organs. – The flower is a more active sink that the leaves. The transformationn of glucose into sucrose is essential for the ability of the flower to use the exogenous glucose. All organs of the floral branch (petals, ovaries and sepals, leaves, stem) are able to form sucrose. The hydrolysis of the obtained sucrose occurs nearly exclusively in the petals.     

Ectopic expression of AINTEGUMENTA in Arabidopsis plants results in increased growth of floral organs.

AINTEGUMENTA (ANT) was previously shown to be involved in floral organ initiation and growth in Arabidopsis. ant flowers have fewer and smaller floral organs and possess ovules that lack integuments and a functional embryo sac. The present work shows that young floral meristems of ant plants are smaller than those in wild type. Failure to initiate the full number of organ primordia in ant flowers may result from insufficient numbers of meristematic cells. The decreased size of ant floral organs appears to be a consequence of decreased cell division within organ primordia. Ectopic expression of ANT under the control of the constitutive 35S promoter results in the development of larger floral organs. The number and shape of these organs is not altered and the size of vegetative organs is normal. Microscopic and molecular analyses indicate that the increased size of 35S::ANT sepals is the result of increased cell division, whereas the increased sizes of 35S::ANT petals, stamens, and carpels are primarily attributable to increased cell expansion. In addition, 35S::ANT ovules often exhibit increased growth of the nucellus and the funiculus. These results suggest that ANT stimulates cell growth in floral organs.     

Floral morphogenesis of Caltha and Trollius (Ranunculaceae) and its systematic significance

The floral morphogenesis of Caltha palustris L. and Trollius buddae Schipcz. was observed with a scanning electron microscope (SEM). The primordia of all floral organs initiate spirally and centripetally and develop centripetally. The spiral initiation sequence may be a basic pattern in Ranunculaceae. The primordia of bracts, sepals, and other floral organs are different in shape: the bract primordia are triangle, the sepal primordia crescent, and the petal (in Trollius), stamen, and carpel primordia hemispheric. This may indicate that the bracts, the sepals and other floral organs are different in origin. The petals are retarded in early developmental stages in Trollius buddae Schipcz, and have purses at the base. The retarded petals are very common in Ranunculaceae and the purse of the petal is similar to that of some Aquilegia species. The microspores in a longitudinal series of stamens develop centripetally in Caltha and Trollius; this may be a basic pattern in Ranunculaceae. The carpel primordia are plicate. In the developmental process of the carpels, the stigmatic tissue appears from the apex of the style and is decurrent along the ventral suture in Caltha, but there is no obvious stigmatic tissue in Trollius. Based on floral morphogenesis characteristics as well as the results from molecular systematics, comparative morphology and palynology studies, we consider that Caltha is not closely related to Trollius and that these two genera should not be treated in the same tribe.     

驴蹄草属和金莲花属(毛茛科)花器官的形态发生及系统学意义

利用扫描电镜观察了驴蹄草Caltha palustris L.和川陕金莲花Trollius buddae Schipcz.花器官的发生和发育过程。结果显示:驴蹄草和川陕金莲花的所有花器官均螺旋状向心式发生、向心式发育,花器官的螺旋状发生方式在毛茛科Ranunculaceae可能是一种基本式样;苞片、萼片与其他花器官原基的形状明显不同,显示苞片、萼片与其他花器官在系统发生上有所不同;川陕金莲花的花瓣在早期延迟发育且基部具囊,花瓣的延迟发育在毛茛科具花瓣的属中非常普遍,而花瓣基部的囊类似于耧斗菜属Aquilegia一些植物;两个属雄蕊群一纵列雄蕊中的小孢子均向心式发育,这种发育方式在毛茛科可能为基本类型。两个属植物的心皮原基均为对折式,在发育过程中,驴蹄草心皮顶端沿腹缝线形成下延的柱头组织,川陕金莲花不形成明显的柱头组织。根据花形态发生和发育特点,并结合其他研究成果,认为这两个属不应当属于同一个族。     

Comparative floral ontogeny in Detarieae (Leguminosae: Caesalpinioideae). 1. Radially symmetrical taxa lacking organ suppression

Flowers in detarioid legume taxa (Isoberlinia angolensis, Microberlinia brazzavillensis, M. bisulcata, Hymenostegia klainii) initiate all 21 floral organs, are radially symmetrical, and have little or no organ suppression. All share a narrow, "Omega"-shaped floral apex and massive bracteoles at initiation. All have helical sepal initiation, starting abaxially. They differ in whether the first sepal initiates medianly (Microberlinia brazzavillensis, M. bisulcata) or nonmedianly (Isoberlinia angolensis, Hymenostegia klainii), and in petal order: helical (I. angolensis) or unidirectional (M. brazzavillensis, M. bisulcata, H. klainii). Stamens initiate in unidirectional order in each whorl except in M. brazzavillensis, which has a bidirectional outer whorl. An unusual feature is the ring meristem in M. bisulcata, on which petals and stamens are initiated. Overlap in time of organ initiation between whorls occurs in I. angolensis, M. brazzavillensis, and M. bisulcata but not in H. klainii. The carpel initiates concurrently with petals in all except H. klainii, in which it initiates with the outer stamens. The carpel remains open at ovule initiation in both species of Microberlinia. These detarioid taxa represent elements of the tribe having essentially radially symmetrical flowers, with all organs initiated and persisting to anthesis, but their specialized "Omega" character-state complex is shared with specialized taxa that have zygomorphic flowers and some organs suppressed.     

Floral ontogeny in Sophoreae (Leguminosae: Papilionoideae): II. Sophora sensu lato (Sophora group)

Floral ontogeny is described in eight species of Sophora sensu lato, representing the Sophora group, as part of a comparative ontogenetic analysis of Polhill's eight groups of tribe Sophoreae, subfamily Papilionoideae. This tribe includes taxa having relatively unspecialized floral structure. Flowers have a five-lobed calyx, a corolla of five free petals, ten mostly unfused, identical stamens, and a carpel. Order of initiation is predominantly acropetal (except for the carpel): sepals, petals, outer stamens plus carpel, inner stamens. Order of initiation within each whorl is unidirectional from the abaxial side. Overlapping initiation among whorls occurs only in S. chrysophylla. Keel petals are slightly fused in six species, and wing petals are fused in 5. tomentosa. Two bird-pollinated species (S. chrysophylla, S. microphylla) lack the papilionaceous corolla of other species, and their petals are unusually long and lack wing sculpturing found in the others. Other floral differences among species mostly involve flower color, differing absolute or relative sizes among organs, and degree of reflexing of vexillum. All but S. davidii have a hypanthium, which develops very late, starting when the bud is about 5 mm long. The distinctions among species (petal size, degree of reflexed position of vexillum, petal sculpturing, color, anther shape, filament hairs, hypanthium presence, calyx lobing) tend to be expressed late in ontogeny.     

Floral Development in Sonja White Clover (Trifolium repens) a Papilionoid Legume

Floral development in Sonja white clover was examined usingscanning electron microscopy. Florets and bracts were foundto arise from common primordia initiated as protuberances fromthe apical meristematic area of the inflorescence. The patternof floret initiation on the inflorescence was acropetal, theoldest florets resting basally. Floral organ initiation withineach floret was acropetal, petals being initiated before stamens.Floret development was zygomorphic, each whorl of floral organsdeveloping unidirectionally from the abaxial side. There wasfound to be overlapping in the timing of initiation and developmentof these organs. Antesepalous stamens were found initially tooutgrow their antepetalous counterparts. Early petal developmentwas synpetalous. Eglandular hairs were found basally on thecalyx cup and on the pedicel. Procumbent hairs were found tobe more numerous and randomly distributed on the abaxial surfacesof the mature calyx cup. Trifolium repens L., Sonja cultivar, white clover, scanning electron microscopy, floral development, inflorescence     

Early flower development in Arabidopsis.

The early development of the flower of Arabidopsis thaliana is described from initiation until the opening of the bud. The morphogenesis, growth rate, and surface structure of floral organs were recorded in detail using scanning electron microscopy. Flower development has been divided into 12 stages using a series of landmark events. Stage 1 begins with the initiation of a floral buttress on the flank of the apical meristem. Stage 2 commences when the flower primordium becomes separate from the meristem. Sepal primordia then arise (stage 3) and grow to overlie the primordium (stage 4). Petal and stamen primordia appear next (stage 5) and are soon enclosed by the sepals (stage 6). During stage 6, petal primordia grow slowly, whereas stamen primordia enlarge more rapidly. Stage 7 begins when the medial stamens become stalked. These soon develop locules (stage 8). A long stage 9 then commences with the petal primordia becoming stalked. During this stage all organs lengthen rapidly. This includes the gynoecium, which commences growth as an open-ended tube during stage 6. When the petals reach the length of the lateral stamens, stage 10 begins. Stigmatic papillae appear soon after (stage 11), and the petals rapidly reach the height of the medial stamens (stage 12). This final stage ends when the 1-millimeter-long bud opens. Under our growing conditions 1.9 buds were initiated per day on average, and they took 13.25 days to progress through the 12 stages from initiation until opening.     

部分小檗科植物的RAPD 分析

利用随机扩增多态性DNA(RAPD)技术分析了部分小檗科(Berberidaceae)5 个属6 种植物:猫儿刺(Berberis julianae Schneid.), 日本绿叶小檗(Berberis thunbergii DC.), 阔叶十大功劳(Mahonia bealei (Fo rt.)Carr.), 南天竹(Nandina domestica Thunb.), 淫羊藿(Epimedium sagittatum(S.et Z.)Maxim.)和八角莲(Dysosma versipellis (Hance)M.Cheng.)。经筛选Sangon 公司的60 个引物, 其中29 个引物的谱带清晰并呈多态性。采用UPGMA 法对求出的遗传距离进行聚类分析, 结果显示:在小檗科内建立十大功劳属(Mahonia)和南天竹属(Nandina)是合理的。     

Developmental programmes in floral organ formation

In contrast to animals, organogenesis in plants is continuous, allowing development in response to intrinsic and extrinsic signals. Organs arise from primordia formed on the flanks of meristems. The apical meristem produces primordia that acquire leaf identity, while floral meristems form primordia which develop into four organ types: sepals, petals, stamens and carpels. The production of mature organs involves two distinct processes, the initiation of organ primordia and the establishment of meristem, primordia and cell identities. Here we concentrate on floral organogenesis in Arabidopsis and examine the extent to which these processes utilize similar control mechanisms and regulatory molecules.