马桑绣球(绣球科)的花器官发生和发育

在扫描电镜下观察了马桑绣球Hydrangea aspera孕性花的发生及发育过程。马桑绣球的花器官向心轮状发生：花萼原基以2/5螺旋式相继发生,花瓣原基几乎同步发生。花瓣开始发育时,与花萼相对的雄蕊发生。与花瓣相对的雄蕊原基与心皮原基几乎同时出现。初始心皮向上扩展,分化出花柱和柱头,向下延伸,嵌入花托,发育为下位子房。花发育成熟时,隔膜于子房的下部连续,而中部和上部不连续,即子房为不完全2室。经过与绣球属已观察过的另外5种1亚种花器官发生和发育比较,发现马桑绣球与藤绣球H. ano mala subs     

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.     

PATTERNS OF FLORAL DEVELOPMENT IN AGALINIS AND ALLIES (SCROPHULARIACEAE). II. FLORAL DEVELOPMENT OF AGALINIS DENSIFLORA

Initiation of floral primordia begins in Agalinis densiflora with production of two lateral adaxial calyx lobe primordia followed by a midadaxial primordium, and then primordia of two abaxial calyx lobes. Initiation of three abaxial corolla lobe primordia is succeeded by that of two stamen pairs and then by primordia of two adaxial corolla lobes. The primordium of the abaxial carpel appears before the adaxial one. Except for the calyx, initiation of primordia proceeds unidirectionally from the abaxial to the adaxial side of the floral apex. Zygomorphy in the calyx, corolla, and androecium is evident during initiation of primordia and is accentuated during organogenesis. The calyx undergoes comparatively rapid organogenesis, but the inner three floral series undergo a protracted period of organogenesis. The perianth series reach maturation prior to meiosis in the anthers. Maturation of the androecium and gynoecium are postmeiotic events.     

Early development of androecia in polystemonous Hydrangeaceae

Polystemonous androecia are diverse in both number and position of stamens. This investigation of polystemonous Hydrangeaceae uses developmental data to characterize (1) the range of developmental variations that account for the diverse androecial patterns and (2) how the expressions of polystemony among Hydrangeaceae compare to those found generally among other angiosperms and especially in their sister family, the Loasaceae, some of which have particularly complex androecia. All polystemonous Hydrangeaceae share the common element of stamen clusters in antesepalous positions. In each of these taxa, the first stamens are initiated opposite the medians of the sepals. Subsequently, stamens form laterally on the flanks of the initial antesepalous stamens, giving rise to the clusters designated as antesepalous triplets. The simplest elaborations based on those common initial developmental steps include (1) adding additional lateral flanking stamens and (2) adding a single stamen in each antepetalous position between adjacent antesepalous groups. More complex elaborations are characteristic of (1) Carpenteria and Philadelphus, which form common primordia at the beginning of androecial development and, subsequently, have stamen primordia form on them, and (2) Deinanthe, which has an elongate hypanthial region on which numerous whorls of stamens are initiated. Carpenteria is unique among Hydrangeaceae in having groups of stamens that are initiated centrifugally in antepetalous positions, and this is similar to complex elements found among some Loasaceae. Generally, the polystemony of Hydrangeaceae that is based in the formation of antesepalous triplets is very similar to that found to evolve in parallel among various clades of rosids and asterids.     

The development of pistillate and perfect florets in Xeranthemum squarrosum (Asteraceae)

The formation of capitulum inflorescence with two different types of floret is an interesting issue in floral biology and evolution. Here we studied the inflorescence, floral ontogeny and development of the everlasting herb, Xeranthemum squarrosum, using epi‐illumination microscopy. The small vegetative apex enlarged and produced involucral bracts with helical phyllotaxy, which subtended floret primordia in the innermost whorl. Initiation of floret primordia was followed by an acropetal sequence, except for pistillate peripheral florets. The origin of receptacular bracts was unusual, as they derived from the floral primordia rather than the receptacular surface. The order of whorl initiation in both disc and pistillate flowers included corolla, androecium and finally calyx, together with the gynoecium. The inception of sepals and stamens occurred in unidirectional order starting from the abaxial side, whereas petals incepted unidirectionally from the adaxial or abaxial side. Substantial differences were observed in flower structure and the development between pistillate and perfect florets. Pistillate florets presented a zygomorphic floral primordium, tetramerous corolla and androecium and two sepal lobes. In these florets, two sepal lobes and four stamen primordia stopped growing, and the ovary developed neither an ovule nor a typical stigma. The results suggest that peripheral pistillate florets in X. squarrosum, which has a bilabiate corolla, could be considered as an intermediate state between ancestral bilabiate florets and the derived ray florets.     

Pseudodiplostemony, and its implications for the evolution of the androecium in the Caryophyllaceae

The androecium of the Caryophyllaceae is varied, ranging from a two-whorled condition to a single stamen. A number of species belonging to the three subfamilies, Caryophyl-loideae, Alsinoideae and Paronychioideae have been studied ontogenetically with the SEM to understand their peculiar androecial development in the broader context of the Caryophyllales alliance. Although patterns of initiation are highly variable among species, there are three ontogenetic modes of stamen initiation: all stamens simultaneous within a whorl, the antepetalous stamens simultaneous and the antesepalous sequentially with a reversed direction, or both whorls sequentially with or without a reversed direction. The most common floral (ontogenetic) sequence of the Caryophyllaceae runs as follows: five sepals (in a 2/5 sequence), the stamens in front of the three inner sepals successively, stamens opposite the two outermost sepals, five antepetalous stamens (simultaneously or in a reversed spiral superimposed on the spiral of the antesepalous stamens), five outer sterile (petaloid) organs arising before, simultaneously or after the antesepalous stamens, often by the division of common primordia. A comparison with the floral configurations of the Phytolaccaceae and Molluginaceae indicates that the outer petaline whorl of the Caryophyllaceae corresponds positionally to the alternisepalous stamens of somePhytolacca, such asP. dodecandra. The difference withP. dodecandra lies in the fact that an extra inner or outer whorl is formed in the Caryophyl-laceae, in alternation with the sepals. A comparable arrangement exists in the Molluginaceae, though the initiation of stamens is centrifugal. A comparison of floral ontogenies and the presence of reduction series in the Caryophyllaceae support the idea that the pentamerous arrangement is derived from a trimerous prototype. Petals correspond to sterillized stamens and are comparable to two stamen pairs opposite the outer sepals and a single stamen alternating with the third and fifth sepals. Petals are often in a state of reduction; they may be confused with staminodes and they often arise from common stamenpetal primordia. The antesepalous stamen whorl represents an amalgamation of two whorls: initiation is reversed with the stamens opposite the fourth and fifth formed sepals arising before the other, while the stamens opposite the first and second formed sepals are frequently reduced or lost. Reductive trends are correlated with the mode of initiation of the androecium, as well as changes in the number of carpels, and affect the antesepalous and antepetalous whorls in different proportions. It is concluded that the androecium of the Caryophyllaceae is pseudodiplos-temonous and is not comparable to diplostemonous forms in the Dilleniidae and Rosidae. The basic floral formula of Caryophyllaceae is as follows: sepals 5—petals 5 (sterile stamens)—antesepalous stamens 3+2—antepetalous stamens 5 gynoecium 5.     

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。     

长角凤仙花（凤仙花科）的花器官发生和发育

以不同发育时期的长角凤仙花Impatiens longicornuta Y.L.Chen（凤仙花科Balsaminaceae）为材料,利用扫描电镜技术观察了其花器官的分化及其发育过程。长角凤仙花为两侧对称花,具2枚侧生萼片,唇瓣囊状,旗瓣具鸡冠状突起,雄蕊5枚,子房上位,5心皮5室。其花器官分化顺序为向心式,萼片—花瓣—雄蕊—雌蕊原基。2枚侧生萼片先发生,然后近轴萼片（即唇瓣）原基和2枚前外侧萼片原基近同时发生;但是这3枚萼片原基的发育不同步,远轴的2枚前外侧萼片原基的发育渐渐滞后,然后停止发育,最后渐渐为周围组织所吸收,直至消失不见。花瓣原基中,旗瓣原基最先发生,4个侧生花瓣原基相继成对发生,且之后在基部成对愈合形成翼瓣;5枚雄蕊原基几乎同时发生,5个心皮原基轮状同时发生。本文结果支持凤仙花属植物为5基数的花,并进一步证实了唇瓣的萼片来源;此外,研究结果表明花器官早期发育资料对植物系统与进化研究具有重要参考价值。     

Elucidating the unusual floral features of Swartzia dipetala (Fabaceae)

In this study, we evaluated the floral ontogeny of Swartzia dipetala, which has peculiar floral features compared with other legumes, such as an entire calyx in the floral bud, a corolla with one or two petals, a dimorphic and polyandrous androecium and a bicarpellate gynoecium. We provide new information on the function of pollen in both stamen morphs and whether both carpels of a flower are able to form fruit. Floral buds, flowers and fruits were processed for observation under light, scanning and transmission electron microscopy and for quantitative analyses. The entire calyx results from the initiation, elongation and fusion of three sepal primordia. A unique petal primordium (or rarely two) is produced on the adaxial side of a ring meristem, which is formed after the initiation of the calyx. The polyandrous and dimorphic androecium also originates from the activity of the ring meristem. It produces three larger stamen primordia on the abaxial side and numerous smaller stamen primordia on the adaxial side. These two types of stamens bear morphologically similar ripening pollen grains. However, prior to the dehiscence of thecae and presentation of pollen in the anther, only the pollen grains of the larger stamens contain amyloplasts. Two carpel primordia are initiated as distinct protuberances, alternating with the larger stamens, in a slightly inner position in the floral meristem, constituting the bicarpellate gynoecium. Both carpels are able to form fruit, although only one fruit is generally produced in a flower. The increase in gynoecium merism probably results in an increase in the surface deposition of pollen grains and consequently in the chance of pollination. This is the first study to thoroughly investigate organogenesis and the ability of the carpel to form fruit in a bicarpellate flower from a member of Fabaceae, in addition to the pollen ultrastructure in the heteromorphic stamens associated with the ‘division of labour’ sensu Darwin. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 303–320.     

Inflorescence and floral development in Astragalus lagopoides Lam. (Leguminosae: Papilionoideae: Galegeae)

Inflorescence and floral organogenesis and development of the bushy perennial legume Astragalus lagopoides of the section Hymenostegis were studied by means of epi-illumination light microscopy. Based on our observations, the primordia of lanceolate racemose inflorescences are born in the axils of leaves. Each inflorescence apex initiates acropetally bracts and floral apices for some time and then eventually ceases meristematic activity and forms an oblong-shaped terminal structure. The formation of such atypical terminal protrusion on the inflorescence meristem is judged to be a diagnostic feature for well-organized cessation of meristem morphogenesis. Pentamerous perfect flowers of the plant show strong zygomorphy and marked overlap in time of initiation among different organ primordia. Unexpectedly, sepal initiation is bidirectional starting from the lateral sides of the floral apex. Other significant developmental feature includes the existence of two types of common primordia, which are formed successively. From the primary common primordia there are produced antesepalous stamens and secondary common primordia. In comparison, the five secondary common primordia subdivide into a petal and an antepetalous stamen primordia. Initiation of two different types of common primordia is possibly the result of rising overlap in time of initiation of organs and demonstrates an advanced developmental style in the genus Astragalus.     

MORPHOLOGICAL STUDIES IN ZYGOPHYLLACEAE. I. THE FLORAL DEVELOPMENT AND VASCULAR ANATOMY OF NITRARIA RETUSA

The floral development and vascular anatomy of Nitraria retusa were investigated in order to understand its characteristic androecium of 15 stamens and to clarify the systematic position of the genus relative to Zygophyllaceae. Sepals arise in a helical sequence and are relatively small at maturity. Petals are initiated almost simultaneously or in a rapid helical sequence. Five stamen primordia arise opposite the sepals. Next, two other antesepalous primordia are incepted centrifugally to the first primordia on the remaining receptacular surface. The outer stamens tend to be squeezed between the petals and upper stamens and appear to make up an antepetalous whorl of stamens. Three carpels arise from a low ringwall and grow into a hairy trilocular pistil. In each locule a single pendulous ovule is present. Disclike nectarial tissue is initiated in pits between the stamens and petals. Long trichomes develop on its surface. It is concluded that the androecium is linked with a haplostemonous condition because the stamens of each triplet develop on strictly localized sectors. The distinction between stamens arising on complex primordia and the inception of three independent units is explained by the “principle of variable proportions.” The vasculature also tends to confirm that the outer stamen pairs belong to antesepalous triplets.     

Floral Ontogeny of Two Species in Magnolia L.

Floral ontogeny is described in two species of genus Magnolia （Magnollaceae）, Magnolia alboserlcea Chun et C. Tsoong, and M. amoena Cheng, representing subgenus Magnolia and subgenus Yulani In Magnolia, by using scanning electron microscope （SEM）. The sequence of Initiation of floral organs Is from proximal to distal. The three distinct outermost and middle organs are Initiated in sequence, but ultimately form a single whorl, thus their ontogeny Is consistent with a sepal Interpretation. The last three tepals （petals） alternate with the preceding tepal whorl. The members of androeclum and gynoeclum arise spirally, although the androecium shows some Intermedlacy between a spiral and whorled arrangement. The carpel prlmordia initiate in group of four to five. The order of stamen Initiation within each tier Is not determined. The floral ontogeny Is remarkably homogeneous between the subgenus Magnolia and subgenus Yulani that does not support the resuming of genus Yulani.     

Floral ontogeny of Ruteae (Rutaceae) and its systematic implications

Floral development was investigated in Ruta graveolens and Psilopeganum sinense, representing two genera in the tribe Ruteae. Special attention was paid to the sequence of initiation of organ whorls in the androecium and gynoecium. The antepetalous stamens arise at the same level as the antesepalous stamens in both species. The carpels are antepetalous in both taxa, indicating the androecium in both genera is obdiplostemonous. Compared with floral ontogeny of the ancestral genus Phellodendron (Toddalioideae), the obdiplostemonous androecium is a derived condition. The floral apex in P. sinense is quadrangular before initiation of the two carpels. Additionally, there are four dorsal and four ventral traces in the ovary. Integrated morphological and anatomical evidence indicates that the bicarpellate gynoecium in Psilopeganum most likely evolved from a tetracarpellate ancestor. Considering the similarities in morphological, geographical and chromosomal features, the ancestor may be Ruta‐like. Further molecular phylogenetic and genetic studies are needed to verify this assumption.     

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.     

Reevaluation of the perianth and androecium in Caryophyllales: implications for flower evolution

The Caryophyllales have the highest diversity in androecial patterns among flowering plants with stamen numbers ranging from 1 up to 4,000. Thanks to the recent progress in reconstructing the phylogeny of core Caryophyllales, questions of floral evolution, such as the origin and diversification of the androecium, can be readdressed. Caryophyllales are unique among core eudicots in sharing an androecial ring meristem or platform with centrifugal development of stamens and petals. Stamens are basically arranged in two whorls and evolution within the clade depends on the shift of either the antesepalous or the alternisepalous whorls to an upper position on the ring meristem and the reduction of the other. Four main developmental phenomena are responsible for the high diversity in androecial patterns: (1) the sterilisation of the outermost stamens through a division of common primordia; (2) the secondary addition of stamens by a centrifugal initiation of supernumerary stamens superimposed on a lower stamen number; (3) the pairwise displacement of alternisepalous stamens to the middle of the outer sepals and their potential fusion, or as part of a pluristaminate androecium; (4) the inversed sequence, reduction and loss of antesepalous stamens. Shifts in stamen numbers depend on pressures of the calyx and carpels and changes in the number of the latter. These patterns are expressed differently in the three main evolutionary lines of core Caryophyllales and are systematically relevant: (1) A basal grade of Caryophyllales, culminating with Caryophyllaceae, Amaranthaceae, Stegnosperma and Limeum, has the antesepalous stamens initiated in upper position on the ring meristem, and alternisepalous stamens are preferentially reduced. Among the antesepalous whorl there is a progressive loss of stamens following a sequence inversed to sepal initiation. Petaloid staminodes are formed by the radial division of outer stamens. (2) The raphide-clade and Molluginaceae are characterized by alternisepalous stamens in upper position on the ring meristem, with a trend to secondary stamen multiplication, and loss of antesepalous stamens. (3) The Portulacineae share the pattern of the raphide clade, but some taxa show shifts to an upper position on the ring meristem of either antesepalous or alternisepalous stamens, linked with secondary multiplications and reduction of either whorl. Different floral characters are plotted on a recent cladogram of Caryophyllales. The data show a consistent correlation between shifting carpel and stamen numbers independent of perianth evolution. Comparative data suggest that the basic androecium of Caryophyllales consists of two whorls of five stamens, linked with an absence of petals, and the evolution of the androecium is a combination of reductions and secondary multiplications of stamens with a highly predictive systematic value.     

Floral ontogeny of Annonaceae: evidence for high variability in floral form

Background and Aims

Annonaceae are one of the largest families of Magnoliales. This study investigates the comparative floral development of 15 species to understand the basis for evolutionary changes in the perianth, androecium and carpels and to provide additional characters for phylogenetic investigation.

Methods

Floral ontogeny of 15 species from 12 genera is examined and described using scanning electron microscopy.

Key Results

Initiation of the three perianth whorls is either helical or unidirectional. Merism is mostly trimerous, occasionally tetramerous and the members of the inner perianth whorl may be missing or are in double position. The androecium and the gynoecium were found to be variable in organ numbers (from highly polymerous to a fixed number, six in the androecium and one or two in the gynoecium). Initiation of the androecium starts invariably with three pairs of stamen primordia along the sides of the hexagonal floral apex. Although inner staminodes were not observed, they were reported in other genera and other families of Magnoliales, except Magnoliaceae and Myristicaceae. Initiation of further organs is centripetal. Androecia with relatively low stamen numbers have a whorled phyllotaxis throughout, while phyllotaxis becomes irregular with higher stamen numbers. The limits between stamens and carpels are unstable and carpels continue the sequence of stamens with a similar variability.

Conclusions

It was found that merism of flowers is often variable in some species with fluctuations between trimery and tetramery. Doubling of inner perianth parts is caused by (unequal) splitting of primordia, contrary to the androecium, and is independent of changes of merism. Derived features, such as a variable merism, absence of the inner perianth and inner staminodes, fixed numbers of stamen and carpels, and capitate or elongate styles are distributed in different clades and evolved independently. The evolution of the androecium is discussed in the context of basal angiosperms: paired outer stamens are the consequence of the transition between the larger perianth parts and much smaller stamens, and not the result of splitting. An increase in stamen number is correlated with their smaller size at initiation, while limits between stamens and carpels are unclear with easy transitions of one organ type into another in some genera, or the complete replacement of carpels by stamens in unisexual flowers.     

台闽苣苔（苦苣苔科）花部器官的形态发生

在扫描电镜下对台闽苣苔 (T .oldhamii (Hemsl.)Solereder)进行了花部器官形态发生的观察 ,为探索该类群的个体发育、类群间的系统发育关系和进化趋势提供依据。研究发现该属植物萼片、花冠和雄蕊发生式样均为五数花类型 ,它们各自来源于花原基上分化出来的萼片原基、花冠原基和雄蕊原基 ;花冠与雄蕊的两侧对称性与花冠上唇生长稍快和退化雄蕊原基发育迟滞相关 ;萼片原基的发生和发育的顺序是不一致的 :萼片原基发生的式样为近轴中原基—远轴 2原基— 2侧原基 ,发育式样则为近轴中萼片— 2侧萼片—远轴 2萼片 ,花蕾时为镊合状排列。花冠裂片原基的发生和发育式样是一致的 ,即远轴中裂原基 (下唇中裂片 )—远轴 2侧裂原基 (下唇 2侧裂片 )—近轴 2裂原基 (上唇 2裂片 )。花蕾期卷迭式为覆瓦状排列 ,从外向内 :下唇中裂片—下唇 2侧裂片—上唇 2裂片或下唇 2侧裂片—上唇 2裂片—下唇中裂片。雄蕊原基与花冠裂片原基互生 ,前方雄蕊原基在发生上稍迟于后方雄蕊原基 ,后者与退化雄蕊原基几乎同时发生 ,但较小 ,并与近轴心皮 (或柱头上唇 )对生。将该属与玄参科 (Scrophulari aceae)的地黄属 (Rehmannia)、苦苣苔科 (Gesneriaceae)的异叶苣苔属 (Whytockia)和尖舌苣苔属 (Rhynchoglossum)的花部器官比较发现     

Comparative ontogeny of perfect and pistillate florets in Senecio vernalis (Asteraceae)

We studied the inflorescence, and in particular ontogeny and development of the florets in Senecio vernalis as a representative member of Asteraceae, using epi-illumination microscopy. Initiation and subsequent development of florets on the highly convex inflorescence apex occur acropetally, except for pistillate ray florets, which show a lag in initiation. Receptacular bracts derive from the receptacular surface after development of all florets. The order of whorl initiation in both disc and ray florets include corolla, androecium and finally the pappus, together with the gynoecium. Development of corolla lobes from a ring meristem occurs in bidirectional order starting from the lateral side, whereas stamens incept unidirectionally from the abaxial side. Concurrently with the inception of two median carpel primordia, a ring meristem develops at the base of the corolla from which pappus bristles differentiate in later stages. Pistillate ray florets show significant differences from perfect disc florets as reflected by the zygomorphic shape of the floral apex and a shift of floral merosity from pentamery to tetramery. Loss of stamens in ray florets occurs due to abortion of primordia after initiation.     

Ontogeny and morphology of the fertile flowers of Hydrangea and allied genera of tribe Hydrangeeae (Hydrangeaceae)

The monophyletic Hydrangeeae (Hydrangeaceae) consists of the clade Cardiandra + Deinanthe and its sister, the Hydrangea clade, which includes the paraphyletic Hydrangea as well as Broussaisia, Decumaria, Dichroa, Pileostegia, Platycrater and Schizophragma. The plesiomorphic imbricate corolla aestivation and polystemony of Cardiandra and Deinanthe distinguish these two genera from most members of the Hydrangea clade. Deinanthe has postgenitally fused styles and cushion-like, dorsally positioned stigmas. The Hydrangea clade is notable because most species of Hydrangea share a floral morphology characterized by small size; tetramerous-pentamerous perianths; inconspicuous sepals; reflexed petals; diplostemony; stamens that are longer than the styles; completely inferior ovaries; separate styles; terminal, papillate stigmas; and dimerous-tetramerous gynoecia. This suite of states is termed the 'Hydrangea floral syndrome' (HFS). Various members of the Hydrangea clade lack the HFS, including (1) Platycrater; (2) Hydrangea anomala; (3) H. paniculata + H. heteromalla; (4) the Schizophragma clade (Schizophragma, Pileostegia and Decumaria); and (5) the macrophylla clade (H. macrophylla, H. scandens, H. hirta, Dichroa and Broussaisia). The meristic uniqueness of Decumaria reflects mutations observed in Arabidopsis (clavata) and Lycopersicon (fasciated) that cause organ number increases because of changes in meristem capital. The modification of early perianth development to form a prominent corolline torus at a point when sepals are diminutive is present in H. anomala and Hydrangea section Cornidia and may be synapomorphic for them. Various transformations in the perianth, androecium, and gynoecium lie behind the floral diversity of Hydrangeeae. Some morphological transformations have been homoplastic, including shifts to polystemony, calyptrate corollas, and synstyly.