ORGANOGENESIS IN THE CARPELLATE FLOWER OF DRIMYS LANCEOLATA

This study of floral development in Drimys lanceolata in Section Tasmannia provides a basis for comparison with D. winteri, a member of the section Wintera, which has been described previously. The carpellate flowers of D. lanceolata have 2 sepals, 4–6 petals, and a solitary carpel, which form in acropetal succession. In symmetry the flower and its apical meristem are bilateral rather than radial, as in the flower of Drimys winteri. The floral apex of D. lanceolata is zonate while that of D. winteri is organized as a mantle and core. Preceding carpel initiation the floral apex of D. lanceolata is narrowly wedge-shaped, while that of D. winteri is low-convex. The entire apex is utilized in carpel initiation in D. lanceolata, involving many subsurface cell divisions over the entire summit. No apical residuum remains, and the carpel is terminal. In this feature the contrast with D. winteri is particularly marked, since in the latter, carpels are initiated laterally around the floral apex, which c an be recognized as an apical residuum after all appendages have formed.     

Original Paper Floral Organogenesis in Verbenaceae sensulato

The early floral ontogeny of three subfamilies, viz. Verbenoideae, Viticoideae and Caryopteridoideae of Verbenaceae (s.l.), was compared. Two differently initiated patterns were found. In the present species of Verbenoideae, there is a unidirectional sequence of organogenesis, from abaxial to adaxial side of the floral apex. While the abaxial paired sepal, petal and stamen arise sequentially, the adaxial paired sepal, petal and stamen do not appear or appear in a much earlier stage. The centripetal whorled sequence of organogenesis appears in Viticoideae and Caryopteridoideae, where sepal primordia arise simultaneously or successively (from adaxial to abaxial). After completion of sepal initiation a plastochron is indicated, during which time a change to the induction of petal takes place, and five petals appear simultaneously, followed by initiation of four stamens. Events of floral organogenesis support the phylogeny inferred from morphological data and rbcL sequence analysis, i.e. the subfamily Verbenoideae does not form a monophyletic group with the subfamilies Viticoideae and Caryopteridoideae.     

Fruit production and floral traits: correlated evolution in <Emphasis Type="Italic">Govenia</Emphasis> (Orchidaceae)

The most common estimate of reproductive success in orchids is usually fruit set. Factors such as resource limitation and certain floral traits may influence reproductive success in animal-pollinated plants. Correlated evolution of reproductive success vs. seven floral traits (inflorescence length, flower number, flower distribution along inflorescence, dorsal sepal length, lateral sepal length, flower color, and column position) was studied in eight species of Govenia. Taxa represented three lineages in the genus. Independent contrasts were calculated on a phylogeny inferred from chloroplast (trnL-F IGS) DNA sequences, and a correlation test and multiple regression were then performed. Two data sets were evaluated, one including all eight species and another excluding G. utriculata, which is autogamous. The historical analyses showed that there is a correlation between reproductive success and dorsal sepal length, column position, and flower number, these correlations suggest that changes in these floral traits usually accompany evolutionary shifts in reproductive success. Multiple regression tests suggest that changes in reproductive success can be explained by shifts in flower number, inflorescence length, column position and by dorsal sepal length. When phylogeny is taken into account, our analyses showed that evolutionary shifts in these floral traits were correlated with changes in reproductive success. Evolutionary correlation between reproductive success and floral traits might be explained by the natural selection of certain floral phenotypes by pollinators.     

灌木铁线莲（毛茛科）花器官的发生与发育

用扫描电子显微镜（SEM）对铁线莲属（Clematis L.）植物灌木铁线莲(C. fruticosa Turcz.)花的形态发生和发育过程进行了观察。灌木铁线莲花原基形成后,4枚萼片以交互对生的方式首先发生,呈轮状排列。最早的4枚雄蕊原基在4枚萼片交接的位置上近螺旋状发生,此后,随着雄蕊原基的向心发生和数目不断增多,其发生的螺旋状序列逐渐明显。雄蕊原基发生后,在花原基顶端,心皮原基沿着雄蕊原基的发生序列呈螺旋状发生。本文结果支持在原始被子植物花中螺旋状排列和轮状排列同时存在的观点。此外,本文也进一步证实了花萼与苞片的同源性。     

马先蒿属花冠无喙类的花器官发生

对花冠无喙类密穗马先蒿(Pedicularis densispica)和大王马先蒿(P．rex)的花器官电镜扫描发现,两种不同花冠型(无齿和具齿)的马先蒿花部器官发生和发育初期十分相似,表现为明显的单轴对称。2个萼片原基首先发生于花顶的近轴侧位,然后沿花顶边缘向远轴端发育形成--马蹄形结构。密穗马先蒿在近轴中部又出现1枚萼片原基,随后马蹄形结构分化出4枚萼片,并与近轴中部的原基愈合后构成5齿萼片;而大王马先蒿的2齿萼片直接由马蹄形结构发育而成。5枚独立的花瓣原基随后发生,但发育相对滞后;除近轴中部位置1枚空缺外,4枚雄蕊原基与花瓣原基位置呈交互发生;2个心皮原基同时在拱形花顶的近轴和远轴端发生,剩余的花顶形成中间的隔膜,并与2个心皮形成中轴胎座。对马先蒿与金鱼草(Antirrhinum majus)和毛地黄(Digitalis purpurea)花器官发生和发育初期的特征进行了比较,讨论了马先蒿属花冠对称性变化的意义。     

Trends in evolution of floral ontogeny in Cassia sensu stricto,Senna, and Chamaecrista (Leguminosae: Caesalpinioideae: Cassieae: Cassiinae); a study in convergence

Distinctions in floral ontogeny among three segregate genera (Cassia sensu stricto, Chamaecrista, and Senna) of Cassia L. support their separation. In all species studied, the order of floral organ initiation is: sepals, petals, antesepalous stamens plus carpel, and lastly antepetalous stamens. Sepal initiation is helical in all three genera, which however differ in whether the first sepal is initiated in median abaxial position (Senna), or abaxial and off-median (Cassia javanica), a rare character state among legumes. Order of petal initiation varies: helical in Senna vs. unidirectional in Cassia and Chamaecrista. Both stamen whorls are uniformly unidirectional. Intergeneric ontogenetic differences occur in phyllotaxy, inflorescence architecture, bracteole formation, overlap of initiation among organ whorls (calyx/corolla in Cassia; two stamen whorls in Chamaecrista), eccentric initiation on one side of a flower, anther attachment, anther pore structure, and precocious carpel initiation in Senna. The asymmetric corolla and androecium in Chamaecrista arise by precocious organ initiation on one side (left or right). The poricidal anther character can result from differing developmental pathways: lateral slits vs. sealing of lateral sutures; clasping hairs vs. sutural ridges; terminal pores (one or two) vs. none; and clamp layer formation internally that prevents lateral dehiscence. Genera differ in corolla aestivation patterns and in stigma type. Convergence is shown among the three genera, based on intergeneric dissimilarities in early floral ontogeny (floral position in the inflorescence, bracteole presence, position of the first sepal initiated, order of petal initiation, asymmetric initiation, overlap between whorls, anther morphology, and time of carpel initiation) resulting in similarities at anthesis (showy, mostly yellow salverform flowers, heteromorphic stamens, poricidal anther dehiscence, bee pollination, and chambered stigma).     

Pentamerous flowers in the genus Phytolacca have been derived from trimerous flowers—New evidence from the floral organogenesis of Phytolacca dodecandra

The floral organogenesis of Phytolacca dodecandra L′Her. (Phytolaccaceae) has been observed under both scanning electron microscope (SEM) and light microscope. The primordia of the floral appendage are arranged according to a pentamerous pattern and acropetal succession. Five sepal primordia arise in a 2/5 sequence, and no petal primordia have been observed. The stamen primordia arise centrifugally. The first two pairs arise successively opposite sepal one and two. In the subsequent initiation of inner and outer stamens, P. dodecandra differs from other species in the genus Phytolacca. The four or five carpel primordia arise in rapid succession, usually equal in number and alternating with the inner stamens. The effects of temporal and spatial factors during the floral organogenesis of P. dodecandra are discussed. The data on the androecial ontogeny in P. dodecandra refute the existence of diplostemony in Phytolaccaceae, in which P. dodecandra occupies a pivotal systematic position. The androecial ontogeny in P. dodecandra supports the viewpoint that in the genus Phytolacca pentamerous flowers have been derived from trimerous flowers.     

ORGAN INITIATION AND DEVELOPMENT OF INFLORESCENCES AND FLOWERS OF ACACIA BAILEYANA

Inflorescence and floral ontogeny are described in the mimosoid Acacia baileyana F. Muell., using scanning electron microscopy and light microscopy. The panicle includes first-order and second-order inflorescences. The first-order inflorescence meristem produces first-order bracts in acropetal order; these bracts each subtend a second-order inflorescence meristem, commonly called a head. Each second-order inflorescence meristem initiates an acropetally sequential series of second-order bracts. After all bracts are formed, their subtended floral meristems are initiated synchronously. The sepals and petals of the radially symmetrical flowers are arranged in alternating pentamerous whorls. There are 30–40 stamens and a unicarpellate gynoecium. In most flowers, the sepals are initiated helically, with the first-formed sepal varying in position. Petal primordia are initiated simultaneously, alternate to the sepals. Three to five individual stamen primordia are initiated in each of five altemipetalous sectorial clusters. Additional stamen primordia are initiated between adjacent clusters, followed by other stamens initiated basipetally as well as centripetally. The apical configuration shifts from a tunica-corpus cellular arrangement before organogenesis to a mantle-core arrangement at sepal initiation. All floral organs are initiated by periclinal divisions of the subsurface mantle cells. The receptacle expands radially by numerous anticlinal divisions in the mantle at the summit, concurrently with proliferation of stamen primordia. The carpel primordium develops in terminal position by conversion of the floral apex.     

Evolutionary loss of sepals and/or petals in detarioid legume taxa Aphanocalyx, Brachystegia, and Monopetalanthus (Leguminosae: Caesalpinioideae)

Floral development using scanning electron microscopy is compared in several taxa of the Brachystegia subtribal group of caesalpinioid tribe Detariae. This group is characterized by missing sepals and/or petals. In Aphanocalyx djumaensis, Monopetalanthus durandii, and two Brachystegia species, one sepal is initiated in median abaxial position. In the first two, one or two additional sepal rudiments may initiate late. Brachystegia species have all five sepals, which remain scalelike. In Aphanocalyx and Monopetalanthus, one petal initiates adaxially and medianly (a position atypical for the first initiated petal in the family); additional petal rudiments may form in lateral sites. In Brachystegia, five petals are initiated unidirectionally on a meristem ring, but all are suppressed after initiation. In all taxa, ten stamens are initiated on a ring meristem: unidirectionally in Monopetalanthus, bidirectionally in Brachystegia, vs. in erratic order in Aphanocalyx. Carpel and petal initiation are concurrent. Different organ whorls overlap in time in Monopetalanthus and Brachystegia. In all, the floral apex characteristically is elongate radially and narrow tangentially after bracteole initiation. Two ontogenetic features, the meristem ring and the radially elongate post-bracteole floral apex, appear to be possible synapomorphies for the Brachystegia group.     

Floral development in Scutellaria pinnatifida (Lamiaceae): the ontogenetic basis for sepal reduction

The Scutellaria is a Labiatae genus (subfamily Scutellarioideae) with a highly specialised floral structure. The genus is characterised by a peculiar two‐lobed calyx with a projecting appendage, named the scutellum. Here, we present a detailed analysis of floral development, using epi‐illumination light microscopy, to clarify open questions about its floral organisation. Floral whorls appeared in an acropetal sequence, with a marked temporal overlap of petal and stamen appearance. Organ appearance in each whorl proceeded unidirectionally from the abaxial to the adaxial side. Significant developmental features included the formation of the scutellum, reduction of sepal lobes and formation of a three‐lobed nectary disc. Our study revealed that both loss of organ initiation and fusion of primordia are responsible for the reduction in sepal members in Scutellaria. The nectary structure was markedly different from most other studied Lamiaceae.     

A morphological and quantitative characterization of early floral development in apple (Malus x domestica Borkh.)

Apple is an important crop and a focus of research worldwide. However, some aspects of floral commitment and morphogenesis remain unclear. A detailed characterization of bourse shoot apex development was undertaken to provide a framework for future genetic, molecular and physiological studies. Eight morphologically distinct stages of shoot apex development, prior to winter dormancy, were defined. Based on measurements of meristem diameter, two stages of vegetative development were recognized. Vegetative meristems were flat, and either narrow (stage 0) or broad (stage 1). Pronounced doming of the apex marked stage 2. During stage 3, the domed meristem initiated four to six lateral floral meristems and subtending bracts before converting to a terminal floral meristem (stage 4). The terminal floral meristem proceeded directly with bractlet and sepal initiation, while lateral floral meristems initiated bractlets (stage 5). Sepal initiation began on the basal lateral flower (stage 6) and continued in an acropetal direction until all floral meristems had completed sepal initiation (stage 7). In this study, only stage 0 and stage 7 apices were observed in dormant buds, indicating that stages 1-6 are transient. The results suggest that broadening of the apex (stage 1) is the first morphological sign of commitment to flowering.     

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.     

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。     

New aspects in floral development of Papilionoideae: initiated but suppressed bracteoles and variable initiation of sepals

BACKGROUND AND AIMS: The increase of molecular data and the resulting insights into legume systematics make the search for new morphological characters and a careful re-investigation of already stated characters necessary. Bracteoles are small, reduced leaves borne close to the base of lateral branches. Although they seem unimportant in older buds, they have an ecological function in protecting the sepal primordia. Furthermore, a morphogenetic function in mediating the onset of sepal initiation is suspected in the literature. The occurrence of bracteoles varies within Papilionoideae, and their distribution is used in legume systematics. But this is open to criticism, because there is a tendency to use 'absent' for 'caducous'. Thus attention here was paid to the initiation of bracteoles as well as to the sequence of sepal initiation. METHODS: The floral development of 30 taxa out of 15 tribes of Papilionoideae was investigated using scanning electron microscopy (SEM). KEY RESULTS: In five taxa the bracteoles initiated, but suppressed early. Furthermore, a broad variability of sepal initiation was found. Besides the widely stated unidirectional pattern, modified unidirectionality, tendencies towards whorled, fully whorled, bidirectional and successive initiation of sepals were all found. CONCLUSION: Initiated but suppressed bracteoles are presented as a 'new' character in Papilionoideae. Considering the presence of bracteoles as a plesiomorphy, their suppression can be seen as a step towards completely reduced bracteoles. The remarkable variability of the sequence of sepal initiation questions the widely stated unidirectionality of organ initiation in Papilionoideae. The different modes of sepal initiation are deducible from the helical pattern of some caesalpinioids, which is seen as a developmental link of the flowers of Papilionoideae and Caesalpinioideae. The bidirectional sepal initiation is possibly a consequence of the morphogenetic function of bracteoles, although bidirectionality is not found in all taxa with reduced bracteoles.     

Floral ontogeny in <Emphasis Type="Italic">Astragalus compactus</Emphasis> (Leguminosae: Papilionoideae: Galegeae): variable occurrence of bracteoles and variable patterns of sepal initiation

Comparative studies of floral ontogeny represent a growing field that promise to provide new insights on floral evolution. Floral ontogenetic information has been used successfully in Leguminosae for re-examining phylogenetic relationships at different levels. Using epi-illumination light microscopy, we present original ontogenetic data in Astragalus compactus, which was chosen because of its unusual arrangement of inflorescence and variable occurrence of bracteoles on flowers. Based on our results, uncommon ontogeny of the inflorescence led to the arrangement of flowers in four different positions. Variation was observed not only in the presence of bracteoles, but also in the order of sepal initiation in flowers of the same inflorescence. Surprisingly, besides the widely stated unidirectional pattern, bidirectional, sequential and an atypical unreported order were observed. High degree of overlapping between whorls and formation of two types of common primordia also were found. The variable occurrence of bracteoles suggests that the species is in an intermediate state towards fully lacking of bracteoles. We propose that the variability of the sequence of sepal initiation is possibly a consequence of the function of mechanical forces generated by surrounding leaves. Relationships between mechanical force and auxin signalling are discussed.     

A floral ontogenetic investigation of the Hydrangeaceae

As a first step towards a broader floral ontogenetic study on Cornales, the flowers of four species of Hydrangeaceae (Deutzia corymbosa, Kirengeshoma palmata, Philadelphus purpurascens and Hydrangea petiolaris) were studied. In Deutzia and Kirengeshoma five sepal primordia are generally initiated, in Philadelphus only four. Sepal initiation in Kirengeshoma shows a tendency to tetramony. Deutzia has a variable sepal initiation. Petal growth was never retarded in the studied species. Initiation of the androecium in Kirengeshoma, Philadelphus and Hydrangea starts with antesepalous primary primordia, on which secondary primordia are soon formed, leading ultimately to the formation of polystaminate androecia. In Deutzia a diplostemonous androecium is formed, starting with the initiation of the antesepalous stamen primordia. Gynoecium development is similar in all species studied: on a concave floral apex, a ring meristem is initiated; it develops into a variable number of continuous carpel primordia, while the centripetally growing common margins form the septa. Initiation of the ovule primordia starts halfway up each placenta and extends in an apical, basal and lateral direction. A number of morphological problems are discussed, such as the derivation of tetramery, and evolutionary and developmental trends in the androecium and gynoecium. Kirengeshoma is well settled in Hydrangeaceae, although its exact position within the family remains uncertain. A sistergroup relationship of Hydrangeaceae with Loasaceae is supported. However, Hydrangeaceae also share features with Saxifragaceae (e.g. similar gynoecium development).     

FLORAL DEVELOPMENT IN CAESALPINIA (LEGUMINOSAE)

Utilizing scanning electron microscopy, we studied the early floral ontogeny of three species of Caesalpinia (Leguminosae: Caesalpinioideae): C. cassioides, C. pulcherrima, and C. vesicaria. Interspecific differences among the three are minor at early and middle stages of floral development. Members of the calyx, corolla, first stamen whorl, and second stamen whorl appear in acropetal order, except that the carpel is present before appearance of the last three inner stamens. Sepals are formed in generally unidirectional succession, beginning with one on the abaxial side next to the subtending bracts, followed by the two lateral sepals and adaxial sepal, then lastly the other adaxial sepal. In one flower of C. vesicaria, sepals were helically initiated. In the calyx, the first-initiated sepal maintains a size advantage over the other four sepals and eventually becomes cucullate, enveloping the remaining parts of the flower. The cucullate abaxial sepal is found in the majority of species of the genus Caesalpinia. Petals, outer stamens, and inner stamens are formed unidirectionally in each whorl from the abaxial to the adaxial sides of the flower. Abaxial stamens are present before the last petals are visible as mounds on the adaxial side, so that the floral apex is engaged in initiation of different categories of floral organs at the same time.     

泽苔草的花器官发生

本文用扫描电镜观察了泽苔草的花器官发生过程,观察结果表明：花萼以螺旋状方式向心发生,花瓣以接近轮状方式近同时发生,不存在花瓣雄蕊复合原基。雄蕊和心皮均以轮状向心方式发生,6枚雄蕊分两轮分别在对萼和对瓣的位置先后发生,至发育的后期排成一轮,但仍分别处于对萼和对瓣的位置;随后发生的第一轮3个心皮原基与3枚萼片相对,第二、三轮心皮原基分别为1~3个,与前一轮心皮相间排列向心发生。本文首次揭示了泽苔草花被的外轮3个萼片螺旋状发生方式,这种螺旋状方式很可能是泽泻科植物的花部结构在进化过程中适应环境而保留下来的一种较原始的叶性特征。     

In vitro culture of young floral buds of tomato (Lycopersicon esculentum mill.)

Young floral buds of tomato (Lycopersicon esculentum Mill.), with sepal primordia only, were cultured successfully in a liquid medium containing Murashige and Skoog's salts, White's vitamins and glycine, 3% (w/v) sucrose and benzylaminopurine (BAP). In 5 weeks, 60–70% of the cultured buds showed normal flower development through to anthesis, and a full complement of floral organs was produced. In the anthers, microsporogenesis occured and microspores and pollen grains were formed. The gynoecium was differentiated into an ovary, style and a stigma, and the ovary contained many ovules, some of which had developing embryo sacs. The floral organs grown in vitro were, however, smaller in size than those produced in vivo. In the control medium, i.e. without BAP, the floral buds produced petal and stamen primordia only and they too showed limited growth. The data presented shows the importance of cytokinins in the initiation and development of organs in the excised floral buds.     

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.