Inflorescence and floral ontogeny in Crocus sativus L. (Iridaceae)

Inflorescence and floral ontogeny of the perennial, herbaceous crop Crocus sativus L. were studied using epi-illumination light microscopy. After production of leaves with helical arrangement a determinate inflorescence forms which becomes completely transformed into a single terminal flower. In some cases, bifurcation of the inflorescence meristem yields two or three floral meristems. The order of floral organs initiation is outer tepals – stamens – inner tepals – carpels. Stamens and outer tepals are produced from the lateral bifurcation of three common stamen-tepal primordia. Within each whorl, organs start developing unidirectionally from the adaxial side, except for the stamens which begin to grow from the abaxial side. Specialized features during organ development include interprimordial growth between tepals forming a perianth tube, fusion at the base of stamen filaments, and formation of an inferior ovary with unfused styles.     

Floral morphogenesis in Euptelea (Eupteleaceae, Ranunculales)

BACKGROUND AND AIMS: Based on molecular phylogenetic studies, the unigeneric family Eupteleaceae has a prominent phylogenetic position at or near the base of Ranunculales, which, in turn, appear at the base of eudicots. The aim of the present paper is to reveal developmental features of the flowers and to put the genus in a morphological context with other basal eudicots. METHODS: Flowers in all developmental stages of Euptelea pleiosperma were collected in the wild at intervals of 7-10 d in the critical stages and studied with a scanning electron microscope. KEY RESULTS: Remnants of a perianth are lacking throughout flower development. Floral symmetry changes from monosymmetric to asymmetric to disymmetric during development. Asymmetry is expressed in that the sequence of stamen initiation is from the centre to both lateral sides on the adaxial side of the flower but starting from one lateral side and proceeding to the other on the abaxial side. Despite the pronounced floral disymmetry, a dimerous pattern of floral organs was not found. The carpel primordia arise between the already large stamens and alternate with them. Stamens and carpels each form a somewhat irregular whorl. The carpels are ascidiate from the beginning. The stigma differentiates as two crests along the ventral slit of the ovary. The few lateral ovules alternate with each other. CONCLUSIONS: Although the flowers have some unusual autapomorphies (wind pollination, lack of a perianth, pronounced disymmetry of the floral base, long connective protrusion, long temporal gap between androecium and gynoecium initiation, small space for carpel initiation), they show some plesiomorphies at the level of basal eudicots (free carpels, basifixed anthers, whorled phyllotaxis), and thus fit well in Ranunculales.     

Floral ontogeny inMazus pumilus (Scrophulariaceae)

InMazus pumilus, all the floral appendages are initiated in acropetal sequence in the second cell layer (except stamens) of the floral primordium by periclinal divisions. The actinomorphic calyx tube is formed due to zonal growth. The zygomorphy in corolla is evident from the inception of petal primordia which arise sequentially as independent units in order of one anterior, a pair of anterio-lateral followed by a pair of posterio-lateral. Later these primordia exhibit differential growth because of which zygomorphy becomes more pronounced. The upper corolla tube is formed by interprimordial growth and lower corolla tube by zonal growth. Stamens are initiated in the third layer of the floral apex. Unlike sepals and petals, in the development of stamens (4) underlying cells of corpus also contribute. Posterior stamen is absent. The stamens become epipetalous because of interprimordial and zonal growth in the common region below the bases of petals as well as stamens. The two carpel primordia arise as crescent shaped structures which become continuous due to interprimordial growth. The ovary is formed by a ring of zonal meristem. The style develops later between stigma and ovary because of intercalary growth. The residual apex grows vertically along with the ovary and forms the septum of the ovary. All the floral appendages exhibit similar pattern of histogenesis and early growth suggesting thereby the appendicular nature of these appendages.     

Ontogeny of Staminate and Carpellate Flowers of Schisandra sphenanthera(Schisandraceae)

The ontogenetic process of the staminate and carpellate flowers of Schisandra sphenanthera Rehd. et Wils., an endemic species to China, was observed for the first time under the scanning electron microscope (SEM). In the staminate flowers, the perianth units and stamens were initiated acropetally in a continuous fasion with 2/5 spiral phyllotaxis, while no female structures were formed. Anthers were differentiated prior to the filaments formation. Throughout all the stages were the stamens arranged spirally on a columniform receptacle. In the carpellate flowers, the initiation sequence of the perianth units and carpels were similar to that of the staminate flowers. In contrast, no male structures were formed. Shortly after initiation, the carpel primordia began their marginal growth besides the apical growth and then appresses were formed on the adaxial surfaces of the primordia. However the lower margins of these appresses were inconspicuous, resulting in conduplicate carpels. Two ovules were developed on the inner surface near either lateral margins of the carpel, shaping laminar placentae. Compared with S. glabra (Brickell) Rehd., a related American species, the evolutionary trend of phyllotaxis of androecia is considered that stamens may change from spiral to approximately whorled arrangement, accompanying with the change of receptacle from a column to a flattened shield. It was also suggested that the stamens being numerous and uncertain in number become certain and decrease in number to 5 (4－7). Sterile stamens are observed and the unisexual nature of the flowers is discussed. Two types of carpel primordia are categorized, corresponding to two types of carpels, namely, ascidiate and conduplicate carpels, respectively.     

Floral Organogenesis and Its Systematic Significance of the Genus Nandina(Berberidaceae)

The initiation of the floral organs of Nandina domestica Thunb. (Berberidaceae) is of the trimemus-whorled pattern. Stamens and petals grew out from the lateral bifurcation of the common stamen-petal primordia; but petals underwent retarded periods of growth in their early development. Carpel initiation belongs to the ascidiate type. Some aspects concerning the trimerous floral organs, the origin of petals, stamen insertion pattem and monocarpellary pistil were discussed. In addition the floral ontogenetic characters among three genera of the Berberidaceae, i.e., Caulophyllum, Podophyllum and Nanclina were compared. It is inferred that the trimerous-whoded arrangement and the diversity of carpel initiation were the two unique characters of Nandina.     

FLORAL DEVELOPMENT IN GYMNOTHECA CHINENSIS (SAURURACEAE)

The development of the inflorescence and flowers are described for Gymnotheca chinensis Decaisne (Saururaceae), which is native only to southeast China. The inflorescence is a short terminal spike of about 50–70 flowers, each subtended by a small bract. There are no showy involucral bracts. The bracts are initiated before the flowers, in acropetal order. Flowers tend to be initiated in whorls of three which alternate with the previous whorl members. No perianth is present. The flower contains six stamens, and four carpels fused in an inferior ovary containing 40–60 ovules on four parietal placentae. Floral symmetry is dorsiventral from inception and throughout organ initiation. Floral organs are initiated in the following order: 1) median adaxial stamen, 2) a pair of lateral common primordia which bifurcate radially to produce two stamen primordia each, 3) median abaxial stamen, 4) a pair of lateral carpel primordia, 5) median adaxial carpel, 6) median abaxial carpel. This order of initiation differs from that of any other Saururaceae previously investigated. The inferior ovary results from intercalary growth below the level of stamen attachment; the style elongates by intercalary growth, and the four stigmas remain free. The floral structure of Gymnotheca is relatively advanced compared to Saururus, but its assemblage of specializations differs from that of either Anemopsis or Houttuynia, the other derived genera in the Saururaceae.     

千金榆花序及花器官发育

在扫描电镜下首次观察了桦木科鹅耳枥属千金榆花序和花的形态发生过程。千金榆雌花序由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化形成2个花原基和2个次级苞片;每个花原基分化出2个心皮原基,形成1个二心皮雌蕊;次级苞片远轴面发育快于近轴面,呈不均等的联合状;雌蕊基部有1层环状花被原基。雄花序为柔荑状,由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化出3个花原基分区,并分化形成3朵小花,小花无花被,位于两侧的小花分别有2枚雄蕊,位于中央的小花有4枚雄蕊,雄蕊共8枚,稀为10枚,该3朵小花为二歧聚伞状排列,其花基数应为2基数。     

ORIGIN AND DEVELOPMENT OF THE TERMINAL CARPEL IN PSEUDOWINTERA TRAVERSII

Flowers of Pseudowintera traversii (Buchan.) Dandy possess a terminal unicarpellate gynoecium. The present study of carpel morphogenesis was initiated for the purposes of (1) providing additional developmental documentation of the occurrence of terminal carpels in the Winteraceae and (2) comparing the mode of initiation and development of the ascidiate terminal carpel of P. traversii with the essentially conduplicate terminal carpel of Drimys lanceolata. Following its axillary origin, the floral apex of P. traversii initiates 2–3 connate sepals, 5–6 petals, 4–15 stamens, and usually a single terminal carpel, in acropetal succession. Bicarpellate gynoecia may occur with a frequency of up to 15 % on a given plant. The floral apex is zonate and shows increased expression of its zonation during later stages of floral development. The terminal carpel is ascidiate from inception and originates as a cylindrical growth around the entire circumference of the floral apex; transformation of the floral meristem into a carpel primordium terminates apical growth of the floral axis. Carpel growth continues to be cylindrical and is mediated by a ring of marginal and submarginal initials at its summit. Earlier and more extensive division of initials and their derivatives on the dorsal rim causes the primordium to become canted adaxially, shifting the apical cleft to a subterminal adaxial position. Continued marginal meristematic activity results in closure of the cleft as well as elevation and elaboration of the stigmatic crests. Five to seven bitegmic ovules are initiated at the same time as crest elaboration and arise in two rows from the adaxial (laminar) position. Carpel maturation is signified by tannin deposition and oil cell differentiation, beginning at the base and proceeding acropetally; carpel margins bordering the cleft are the last to differentiate. Carpel procambialization is continuous and acropetal from inception, with the dorsal median bundle differentiating before the ventral strands. The significance of occasional bicarpellate flowers is discussed.     

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.     

OVERLAPPING ORGAN INITIATION AND COMMON PRIMORDIA IN FLOWERS OF PISUM SATIVUM (LEGUMINOSAE: PAPILIONOIDEAE)

The floral ontogeny of Pisum sativum shows a vertical order of succession of sepals, petals plus carpel, antesepalous stamens, and antepetalous stamens. Within each whorl, unidirectional order is followed among the organs, beginning on the abaxial side of the flower, as in most papilionoids. Unusual features include the four common primordia which precede initiation of discrete petal and antesepalous stamen primordia, and the marked overlap of organ initiations between whorls which are usually separately initiated. The stamens arise in free condition, then become diadelphous by intercalary growth at the base of nine stamens, and finally become pseudomonadelphous by surface fusion between the vexillary stamen filament and the adjacent edges of the filament tube. The early initiation of the carpel is not unique among papilionoids, but is somewhat unusual.     

LOSS OF FLORAL ORGANS IN ATELEIA (LEGUMINOSAE: PAPILIONOIDEAE: SOPHOREAE)

Ateleia herbert-smithii is unique among legumes in being a wind-pollinated tree; carpellate and staminate flowers are restricted to different trees. Development of the two floral morphs, however, is essentially the same except for smaller carpels in functionally staminate flowers and failure of pollen formation in the anthers of functionally carpellate flowers. The floral development of Ateleia herbert-smithii is highly atypical among papilionoids and the tribe Sophoreae. Order of organ initiation is: sepals, solitary petal, carpel, and lastly all stamens in erratic order. Sepal order is unidirectional from the abaxial side, the normal pattern for papilionoids. Only one petal, the vexillum or standard, is initiated. Subsequent initiation is completely different from the usual unidirectional pattern of most papilionoids. A meristem ring forms, delimiting the solitary carpel centrally. Ten stamen primordia are initiated on the meristem ring, first laterally, then adaxially, and lastly abaxially. There is a tendency for antesepalous stamens to form before the antepetalous ones. The loss of four of the five petals is thought to alter drastically the subsequent organogeny as to position of organs and their order of initiation. Carpel initiation in Ateleia is precocious, but not uniquely so among legumes.     

商陆属植物的花器官发生

为进一步研究商陆科的系统位置提供花器官发生和发育的证据,在扫描电子显微镜下观察了商陆Phytolacca acinosa、多雄蕊商陆P. polyandra和垂序商陆P. americana的花器官发生.结果表明: 商陆属植物花被的发生均为2/5型螺旋发生.在同一个种不同的花蕾中,花被的发生有两种顺序:逆时针方向和顺时针方向.远轴侧非正中位的1枚先发生.雄蕊发生于环状分生组织.在单轮雄蕊的种中8-10枚雄蕊为近同时发生;两轮雄蕊的种8枚内轮雄蕊先发生,6-8枚外轮雄蕊随后发生,内轮雄蕊为同时发生,外轮雄蕊发生次序不规则.心皮原基也发生于环状分生组织,8-10枚心皮原基为同时发生.在后来的发育过程中,商陆的心皮发育成近离生心皮雌蕊;其他2种心皮侧壁联合发育成合生心皮雌蕊.对商陆属植物花器官发生的类型及发育形态学做了分析,结果支持商陆科在石竹目系统发育中处于原始地位的观点.     

INFLORESCENCE AND FLORAL DEVELOPMENT IN HOUTTUYNIA CORDATA (SAURURACEAE)

The inflorescence of Houttuynia cordata produces 45–70 sessile bracteate flowers in acropetal succession. The inflorescence apical meristem has a mantle-core configuration and produces “common” or uncommitted primordia, each of which bifurcates to form a floral apex above, a bract primordium below. This pattern of organogenesis is similar to that in another saururaceous plant, Saururus cernuus. Exceptions to this unusual development, however, occur in H. cordata at the beginning of inflorescence activity when four to eight petaloid bract primordia are initiated before the initiation of floral apices in their axils. “Common” primordia also are lacking toward the cessation of inflorescence apical activity in H. cordata when primordia become bracts which may precede the initiation of an axillary floral apex. Many of these last-formed bracts are sterile. The inflorescence terminates with maturation of the meristem as an apical residuum. No terminal flowers or terminal gynoecia were found, although subterminal gynoecia or flowers in subterminal position may overtop the actual apex and obscure it. Individual flowers have a tricarpellate syncarpous gynoecium and three stamens adnate to the carpels; petals and sepals are lacking. The order of succession of organs is: two lateral stamens, median stamen, two lateral carpels, median carpel. The three carpel primordia almost immediately are elevated as part of a gynoecial ring by zonal growth of the receptacle below the attachment of the carpels. The same growth elevates the stamen bases so that they appear adnate to the carpels. The trimerous condition in Houttuynia is the result of paired or solitary initiations rather than trimerous whorls. Symmetry is bilateral and zygomorphic rather than radial. No evidence of spiral arrangement in the flower was found.     

Floral development of Justicia gendarussa (Acanthaceae)

The sequence of primordial initiation is acropetal and the primordia develop in the same order in which they appear. The floral apex has a two-layered tunica in all stages of development. There is no significant difference in the initiation of any of the floral appendages and thus all floral organs are homologous with respect to their histogenetic origin. The short calyx tube is formed by ontogenetic fusion of the bases of sepals; but the corolla tube arises partly by ontogenetic union of originally free parts and partly by zonal or intercalary growth. Each primordium receives a single procambium strand shortly after its initiation, except those of the posterior pair of stamens, which do not receive any vascular supply. This provides another example which goes against the doctrine of "conservatism of vascular bundles". The placentation in Justicia is parietal ontogenetically as well as anatomically.     

Conversion of perianth into reproductive organs by ectopic expression of the tobacco floral homeotic gene NAG1.

Mutations in the AGAMOUS (AG) gene of Arabidopsis thaliana result in the conversion of reproductive organs, stamens and carpels, into perianth organs, sepals and petals. We have isolated and characterized the putative AG gene from Nicotiana tabacum, NAG1, whose deduced protein product shares 73% identical amino acid residues with the Arabidopsis AG gene product. RNA tissue in situ hybridizations show that NAG1 RNA accumulates early in tobacco flower development in the region of the floral meristem that will later give rise to stamens and carpels. Ectopic expression of NAG1 in transgenic tobacco plants results in a conversion of sepals and petals into carpels and stamens, respectively, indicating that NAG1 is sufficient to convert perianth into reproductive floral organs.     

Floral development of dioecious species and trends of floral evolution in Piper sensu lato

The initiation of the floral parts (mainly stamens and carpels) is described for the four dioecious species of Piper: Piper polysyphorum C. DC, P. bavinum C. DC., P. pedicellatum C. DC., P. pubicatulum C. DC. The initiation order resembles that in the perfect flowers of some species, such as P. amalago. The carpels are initiated simultaneously, in most cases, as three primordia. In P. polysyphorum , carpel tips split into two lobes, so that finally a four- or five-lobed stigma will be formed when the ovary is fully developed. The staminodes (exactly, staminodial primordia) in the female flowers are initiated in the same order as the stamens in the male flowers and remain until the ovaries are enclosed. The unisexual flowers have stamens reduced to three or two. The reduction of stamen or staminode (staminodial primordium) number is accompanied by the change of their positions from opposite the carpels to alternate. After the initiation of the staminodes, or, exactly staminodial primordia, in the female flowers, the central part of the floral apex forms a ring meristem which is triangular. The carpel primordia (often three) are initiated on the three points of the ring meristem. The evolutionary trends of the flowers of Piper sensu lato are discussed.     

Flower development inAntirrhinum majus L. (Scrophulariaceae) with a comment upon corolla tube formation

The inflorescence apex is stratified and has a single layered tunica, bracts as well as all the floral organs develop acropetally. Except for stamens which are initiated with the activity of deeper layers of apex, all other floral appendages are initiated with the activity of second layer. On the basis of ontogenetic studies the floral organs have been homologised with the leaves. The adjacent calyx lobes develop and remain closely appressed and do not show post-genital fusion. Corolla tube is formed due to zonal growth preceeded by joint growth of hump on the abaxial face of stamen primordia and interprimordial region between petal primordia. Thus no post-genital fusion was observed. The placentae develop concomitantly with carpellary wall.     

DIOECY IN BAUHINIA RESULTING FROM ORGAN SUPPRESSION

Bauhinia malabarica and B. divaricata have both been reported to have dimorphic flowers; floral development of these species has been investigated and compared using SEM. B. malabarica is subdioecious, with three types of flowers: perfect, staminate, and carpellate. Individual trees usually have only one type of flower. Perfect and carpellate flowers have similar initiation of floral organs; each has five sepals, five petals, two whorls of five stamen primordia and a carpel primordium. The carpels of carpellate flowers do not differ from those of perfect flowers throughout development. Both have a gynophore or stipe and a cuplike hypanthium. Stamen development diverges markedly after mid-development: the perfect flowers have ten stamens in two whorls, the outer with longer filaments than the inner. All stamens have anthers, which are covered abaxially with abundant inflated trichomes. Carpellate flowers have a circle of short cylindrical staminodia, each bearing a few hairs, about the base of the carpel on the rim of the hypanthium. Heteromorphy in B. malabarica is effected by suppression of stamen development, even though the usual number of stamen primordia is initiated. Suppression of stamens occurs at midstage in development in carpellate flowers of B. malabarica, and is complete. In B. divaricata nine stamen primordia are released from suppression in late stage, undergo intercalary growth and form a staminodial tube around the carpel stipe. The dimorphy in B. divaricata is expressed late in bud enlargement as divergent rates of growth in the carpel in the two morphs.