Studies on the Development of Flower in Gastrodia ekata Bl.

The flower of Gastrodia elata B1. grows usually solitary in the bract-axial. The parts of flower are appeared in the following order: sepals, petals, stamens and pestil. In addition, the bases of calyx and corolla unite forming the oblique “floral tube”. Inner three petals, the median lip of the petal is usually larger and different in shape. The lip is three lobes, the marginal and its bases meristem keeps its activity forming lobes marginal tassel, at the base forming one fleshy reniform callus on both sides of lip. A column is united by stamen and style. The stamen is on the top at the back of rostellum in the column. Three stamens primodia of them, only one can developed, on the lateral stamens, during the early stage of flower development, it will not development further and stays on rudimentary, it forming two horn-like protuberances of the clinandrium.     

Genes directing flower development in Arabidopsis.

We describe the effects of four recessive homeotic mutations that specifically disrupt the development of flowers in Arabidopsis thaliana. Each of the recessive mutations affects the outcome of organ development, but not the location of organ primordia. Homeotic transformations observed are as follows. In agamous-1, stamens to petals; in apetala2-1, sepals to leaves and petals to staminoid petals; in apetala3-1, petals to sepals and stamens to carpels; in pistillata-1, petals to sepals. In addition, two of these mutations (ap2-1 and pi-1) result in loss of organs, and ag-1 causes the cells that would ordinarily form the gynoecium to differentiate as a flower. Two of the mutations are temperature-sensitive. Temperature shift experiments indicate that the wild-type AP2 gene product acts at the time of primordium initiation; the AP3 product is active later. It seems that the wild-type alleles of these four genes allow cells to determine their place in the developing flower and thus to differentiate appropriately. We propose that these genes may be involved in setting up or responding to concentric, overlapping fields within the flower primordium.     

FLORAL ANATOMY OF KRAMERIA LANCEOLATA

The anatomy of each of the series of floral organs of Krameria lanceolata was examined. The sepals are characterized by three main veins each, an undifferentiated mesophyll, and stomata on the upper epidermis. The fleshy petals are distinguished by their numerous veins as well as by palisade-like epidermal cells on the outer surface. The three partially united petals have each a single vein and long, narrow epidermal cells similar to those on other floral organs. The stamens are united at their bases and bear tetra-sporangiate, conical anthers. The gynoecium includes a sterile and a fertile carpel. In the receptacle the veins to the sepals and petals are separated by a wide gap; those to the petals and stamens, by a narrow gap. Anatomical characteristics of the flower dissociate Krameriaceae from the legumes with which they have frequently been thought to be allied.     

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.     

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.     

Spatial fragrance patterns within the flowers ofRanunculus acris (Ranunculaceae)

Floral scents emitted from different flower parts ofRanunculus acris were investigated by trapping headspace volatiles onto Porapak Q followed by solvent desorption and GC-MS analysis. Isoprenoids, strongly dominated bytrans--ocimene, constituted the principal class of volatiles in all flower parts except pollen; sesquiterpenes were especially diverse. Odors collected separately from petals, stamens, and sepals + gynoecium comprised the same volatiles, but these were present in disparate proportions among the flower parts, thereby creating subtle contrasts within the flower. The main sources of volatiles were the petals and stamens, which made equal contributions to the whole-flower fragrance. Emissions from the petals differed quantitatively between the apical and basal petal regions, thereby paralleling optical nectar-guide patterns. Pollen odor was markedly unlike that of other flower parts, with only few volatiles, a high representation of 5-methylene-2(5H)-furanone (protoanemonin), and no detectabletrans--ocimene. The distinctiveness of the pollen's volatile profile suggests that it may serve a signalling role to pollen-feeding insects.     

FLORAL DEVELOPMENT IN MANGROVE RHIZOPHORACEAE

The flowers of mangrove Rhizophoraceae (tribe Rhizophoreae) are adapted to three different pollination mechanisms. Floral development of representative species of all four genera suggests that the ancestral flower of the tribe was unspecialized, with successively initiated whorls of separate sepals, petals, antisepalous stamens, and antipetalous stamens; at its inception, the gynoecium had a united, half-inferior ovary and separate stigmatic lobes. This developmental pattern is found in Rhizophora mangle (wind-pollinated) and Ceriops decandra (insect-pollinated). In Kandelia, all floral organs distal to the sepals are initiated simultaneously, and there has apparently been an evolutionary amplification in the number of stamens to about six times the number of petals. Explosive pollen release evolved independently in C. tagal and in Bruguiera. In the former, all stamens belong to one whorl and arise simultaneously upon a very weakly differentiated androecial ring primordium. In Bruguiera, the androecial ring is pronounced, and two whorls of stamens arise upon it; the primordia of the antisepalous whorl arise first but are closer to the center of the apex than the antipetalous stamen primordia. The antisepalous stamens bend toward and are enclosed by the petals early in development. In all genera, the inferior ovary develops by zonal growth of receptacular tissue; additional intercalary growth above the placenta occurs in Bruguiera. In general, floral specialization is accompanied by an increase in the width of the floral apex compared to the size of the primordia, increasing fusion of the stylar primordia, and decreasing prominence of the superior portion of the ovary. Apparent specializations of petal appendages for water storage, including the presence of sub-terminal hydathodes (previously unreported in any angiosperm), were found in two species in which flowers remain open during the day but were absent from two species normally pollinated at night or at dawn. Distinctive tribal characteristics that may aid in phylogenetic analysis include the mode of development of the inferior ovary; the aristate, bifid, usually fringed petals that individually enclose one or more stamens; the intrastaminal floral disc; and the initially subepidermal laticiferous cell layer in the sepals and ovary.     

Phyllotaxis and the Initiation of Primordia During Flower Development in Silene

The measured divergence angles between successive primordiain the developing flower were compared with angles expectedon several hypotheses concerning primordial initiation. Theresults lead to the conclusion that the position and sequenceof initiation of the younger sepals is determined by the olderones but that the influence of an older primordium lasts foronly two plastochrons. The petals and carpels are apparentlypositioned by the sepals. The positions of the stamens are consistentwith their king determined by the sepals (antesepalous stamens)or petals (antepctalous stamens), but their sequence of initiationis consistent with its being determined, like the sepals, bythe two youngest primordia. The data indicate that there aretwo sets of factors governing the initiation of the primordiasubsequent to the sepals: one governing the positioning of theprimordia and resembling the factors governing the positionsof axillary buds, and the other governing the sequence of primordiaand resembling the factors which determine the initiation ofleaves. Measurements of the plastochron ratios were used tocalculate the sizes of the sepal, petal and stamen primordiaat initiation. At the moment of initiation the sepal primordiawere about one third, and the petal and stamen primordia aboutone sixth, of the size of the leaf primordia. In its early developmentthe Silene flower therefore resembles a condensed leafy shootwith precocious axillary buds but with primordia which are smallcompared to leaf primordia. Silene coeli-rosa, flower development, primordia, phyllotaxis     

FLORAL ONTOGENY IN SOPHOREAE (LEGUMINOSAE: PAPILIONOIDEAE). I. MYROXYLON (MYROXYLON GROUP) AND CASTANOSPERMUM (ANGYLOCALYX GROUP)

Floral ontogeny is described and documented using scanning electron microscopy in Myroxylon balsamum and Castanospermum australe, representatives respectively of Polhill's Myroxylon and Angylocalyx groups (Leguminosae: Papilionoideae), groups exhibiting relatively unspecialized flower structure for the tribe Sophoreae. Both are woody tropical trees with axillary or terminal racemes or panicles. Bracteoles are present in both Myroxylon and Castanospermum. Flowers are initiated singly in bract axils, which are produced in acropetal order by the inflorescence apical meristem. The flower structure of both includes a broad calyx tube, five petals lacking any fusion (only the vexillary distinctive), ten free homogeneous stamens in two whorls, and a long-stipitate carpel. The two taxa are alike in early organogenetic stages with essentially acropetal order of initiation: sepals, petals, outer stamens plus carpel, inner stamens. Within each whorl the order is unidirectional from the abaxial side. They are alike through middevelopment with one exception. There is accelerated vexillar enlargement in Castanospermum by middevelopment, not found in Myroxylon. Both have a hypanthium, which forms late in development. In both, large flower size, exserted stamens, and hypanthium are adaptations to bird-pollination. Differences between the two that are manifested in late development include strongly zygomorphic calyx and petal color change over time (Castanospermum), stamens sagittate and apiculate with some basal filament fusion (Myroxylon), stigma form differences, and fruit form.     

Flower development of Meliosma (Sabiaceae): evidence for multiple origins of pentamery in the eudicots

Flower developmental studies are a complement to molecular phylogenetics and a tool to understand the evolution of the angiosperm flower. Buds and mature flowers of Meliosma veitchiorum, M. cuneifolia, and M. dilleniifolia (Sabiaceae) were investigated using scanning electron microscopy to clarify flower developmental patterns and morphology, to understand the origin of the perianth merism, and to discuss the two taxonomic positions proposed for Sabiaceae, among rosids or in the basal grade of eudicots. Flowers in Meliosma appear pentamerous with two of the five sepals and petals strongly reduced, three staminodes alternating with two fertile stamens opposite the small petals, and a two-carpellate gynoecium. The flower development in Meliosma is spiral without distinction between bracteoles and sepals. Because of this development, sepals, petals, and stamens are almost opposite and not alternating as expected in cyclical pentamerous flowers. In four-sepal flowers the direction of petal initiation is reversed. The symmetry of the flower appears to be transversally zygomorphic, although this is hidden by the almost equal size of the larger petals. Evidence points to a unique pentamerous origin of flowers in Meliosma, and not to a trimerous origin, as earlier suggested, and adds support to multiple origins of pentamery in the eudicots.     

THE DIFFERENTIATION OF PIGMENTATION IN FLOWER PARTS. I. THE FLAVONOID PIGMENTS OF IMPATIENS BALSAMINA,GENOTYPE IIHHPrPr,AND THEIR DISTRIBUTION WITHIN THE PLANT

Chromatographic analysis of stems, sepals and petals of inbred Impatiens balsamina of the red-flowered genotype llHHP^rP^r has revealed a characteristic assemblage of flavonoid pigments in each organ. The more conspicuous compounds have been identified or partially characterized. The stems possess 3-monoglucosides of kaempferol, quercetin, pelargonidin, cyanidin and, presumably, delphinidin. The variety of pigments is less in flower parts than in stems, and less in petals than in sepals, but the flower parts exhibit a greater elaboration of substituents on the aromatic nuclei. The paired petals of mature flowers are pigmented by p-coumaroyl and feruloyl esters of pelargonidin-3, 5-diglucoside supplemented by more highly substituted derivatives of pelargonidin and by large amounts of kaempferol as the aglycone and two glucosides. The distribution of pigments has significance in the biology of the plant as well as providing an approach to studies of factors which control flower differentiation.     

Structure and development of ‘witches' broom’ galls in reproductive organs of Byrsonima sericea (Malpighiaceae) and their effects on host plants

Galls are anomalies in plant development of parasitic origin that affect the cellular differentiation or growth and represent a remarkable plant–parasite interaction. Byrsonima sericea DC. (Malpighiaceae) is a super host of several different types of gall in both vegetative and reproductive organs. The existence of galls in reproductive organs and their effects on the host plant are seldom described in the literature. In this paper, we present a novel study of galls in plants of the Neotropical region: the ‘witches' broom’ galls developed in floral structures of B. sericea. The unaffected inflorescences are characterised by a single indeterminate main axis with spirally arranged flower buds. The flower buds developed five unaffected brownish hairy sepals and five pairs of elliptical yellow elaiophores, five yellow fringed petals, 10 stamens and a pistil with superior tricarpellar and trilocular ovary. The affected inflorescences showed changes in architecture, with branches arising from the main axis and flower buds. The flower buds exhibited several morphological and anatomical changes. The sepals, petals and carpels converted into leaf‐like structures after differentiation. Stamens exhibited degeneration of the sporogenous tissue and structures containing hyphae and spores. The gynoecium did not develop, forming a central meristematic region, from which emerges the new inflorescence. In this work, we discuss the several changes in development of reproductive structures caused by witches' broom galls and their effects on reproductive success of the host plants.     

Type specification and spatial pattern formation of floral organs: A dynamic development model

A mathematical model simulating spatial pattern formation (positioning) of floral organs is proposed. Computer experiment with this model demonstrated the following sequence of spatial pattern formation in a typical cruciferous flower: medial sepals, carpels, lateral sepals, long stamens, petals, and short stamens. The positioning was acropetal for the perianth organs and basipetal for the stamens and carpels. Organ type specification and positioning proceed non-simultaneously in different floral parts and organ type specification goes ahead of organ spatial pattern formation. Computer simulation of flower development in several mutants demonstrated that the AG and AP2 genes determine both organ type specification and formation of the zones for future organ development. The function of the AG gene is to determine the basipetal patterning zones for the development of the reproductive organs, while the AP2 gene maintains proliferative activity of the meristem establishing the acropetal patterning zone for the development of the perianth organs.     

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.     

The homeotic Macho mutant of Antirrhinum majus reverts to wild-type or mutates to the homeotic plena phenotype

Plants of Antirrhinum majus carrying the semidominant Macho alleles of the plena gene display carpelloid sepals and staminoid petals, but the two inner flower whorls of stamens and carpels are normal and produce fertile gametes. In the recessive plena mutant, in contrast, the two outer whorls are normal whereas the stamens are largely or entirely petaloid and the carpels sepaloid, thus producing weakly male-fertile or fully sterile lines. Two new plena and two new Macho alleles have been induced in transposon tagging experiments. Genetic and molecular analysis revealed that the two contrasting mutant phenotypes are caused by mutations in one and the same gene: Several wild-type plants appeared among 27 000 F1 plants of a cross between Macho female plants and wild-type males bearing the active transposons Taml and Tam3. One of these plants segregated plena mutants, three showed reversions to wild-type and another two segregated Macho plants, possibly representing somatic reversions. Additional evidence was provided by an allelism test of Macho × plena. Molecular analysis has independently corroborated the genetical results. Moreover, the double mutant Macho/deficiens shows only carpels and plena/deficiens only sepals, which is in accord with combinatorial models for homeotic flower formation presented recently.     

The Arabidopsis thaliana flower organ specification gene regulatory network determines a robust differentiation process

The Arabidopsis thaliana flower organ specification gene regulatory network (FOS-GRN) has been modeled previously as a discrete dynamical system, recovering as steady states configurations that match the genetic profiles described in primordial cells of inflorescence, sepals, petals, stamens and carpels during early flower development. In this study, we first update the FOS-GRN by adding interactions and modifying some rules according to new experimental data. A discrete model of this updated version of the network has a dynamical behavior identical to previous versions, under both wild type and mutant conditions, thus confirming its robustness. Then, we develop a continuous version of the FOS-GRN using a new methodology that builds upon previous proposals. The fixed point attractors of the discrete system are all observed in the continuous model, but the latter also contains new steady states that might correspond to genetic activation states present briefly during the early phases of flower development. We show that both the discrete and the continuous models recover the observed stable gene configurations observed in the inflorescence meristem, as well as the primordial cells of sepals, petals, stamens and carpels. Additionally, both models are subjected to perturbations in order to establish the nature of additional signals that may suffice to determine the experimentally observed order of appearance of floral organs. Our results thus describe a possible mechanism by which the network canalizes molecular signals and/or noise, thus conferring robustness to the differentiation process.     

太原黄耆(豆科)花器官及胚珠发育研究

太原黄耆是新近发表的物种,分布于中国陕西和山西。该实验利用扫描电子显微镜对太原黄耆的花器官发生和发育过程进行观察研究。结果显示:(1)太原黄耆的各轮花器官都是从远轴端向近轴端单向连续发生,在不同轮之间存在花器官重叠发生的现象。(2)在花的发育过程中,出现2种共同原基,即初级共同原基和次级共同原基,由初级共同原基发育成对萼雄蕊原基和次级共同原基,再由次级共同原基发育成花瓣原基和对瓣雄蕊原基。(3)雄蕊管近轴端基部开口是在进化过程中产生的特殊结构,是一种对传粉者的适应机制,从而有利于传粉活动的进行。(4)胚珠为倒生胚珠,具有2层珠被,认为倒生胚珠是内外2层珠被共同作用的结果。     

A new species of Blakea (Melastomataceae) from Panama with foliaceous sepal appendages and zygomorphic flowers

The new species Blakea bocatorena is described from Bocas del Toro province in Panama. It is only the third species to be described in the genus that has foliaceous appendages on the sepals. The other two species with these sepal appendages are B. calycosa and B. tuberculata. Blakea bocatorena differs from the latter two species in that it has white petals and a type of herkogamy in which the style is opposite the stamens and results in a zygomorphic flower. In B. calycosa and B. tuberculata, the larger petals are pink-magenta and tuberculate with the stamens encircling the exserted style resulting in a radially symmetric herkogamous flower.     

Continuous petal variation in Epimedium tianmenshanense (Berberidaceae), a taxon endemic to western Hunan,China

In the protologue, Epimedium tianmenshanense (Berberidaceae), a species endemic to western Hunan, China, was described as ‘flower small, 0.2–0.4 cm diam., inner sepals white, petals as long as inner sepals or a little shorter than the latter, spur very short, ca 5.0 mm’. However, both morphological characteristics and molecular evidence suggest that E. tianmenshanense is closely related to E. baojingense, a taxon with a long spur, thus suggesting that the size of the floral parts is not as reliable as previously believed. When investigating the variability of E. tianmenshanense in more detail, in the field as well as in cultivation, we found that the petals were are highly variable in morphology (both shape and size), being cucullate, subulate, short to long spurred, and with various transitions. The flowers size varied from small to large accordingly. The flowers with cucullate and subulate petals, which were a little shorter than the inner sepals or almost as long as the latter, were small (about 0.8 cm in diameter). The flowers with long spurs, which were much longer than the inner sepals, were also large (about 2.5–3.5 cm in diameter). Finally, the flowers with short spurs, which were a little longer than the inner sepals, were medium-sized (about 1.0–1.2 cm in diameter). In addition, the color of inner sepals was revised from ‘white, occasionally light mulberry-purple’ to yellowish green or yellowish white. Epimedium tianmenshanense is a perfect example of natural petal evolution, which could be used for further taxonomic and evolutionary studies. The reason for the variation and the taxonomic treatment of the species still need further study.     

STUDIES IN THE FLORAL ANATOMY OF POLYGALA (POLYGALACEAE)

Flowers of Polygala are irregular, each having three small and two long winglike sepals, three petals, eight stamens, and a bilocular ovary. These flowers have been considered pentamerous, and placentation has been subject to various interpretations. Development and anatomy of flowers of Polygala alba, P. lanceolata, and P. lutea were studied to see if evidence of pentamery and change in placentation could be found. These studies reveal no evidence of vestigial petals or stamens nor of vascular traces to organs that are missing in the three species studied. Neither are there abortive sporangia in the bisporangiate anthers. Observations on development of carpel primordia and on the vascular plan of mature carpels indicate that placentation is fundamentally parietal rather than axile. Speculation is offered as to the derivation of this type of placentation.