Floral organogenesis of Delavaya toxocarpa （Sapindaceae; Sapindales）

The floral organogenesis and development of Delavaya toxocarpa Franch. （Sapindaceae） were studied trader scanning electron microscope and light microscope to determine its systematic position within Sapindaceae. Flowers arise in terminal thyrses. The sepal primordia initiate in a spiral （2/5） sequence, which are not synchronous. The five petal primordia initiate almost synchronously and alternate with sepal primordia. Eight stamens initiate almost simultaneously and their differentiation precedes that of the petals. The last formed petal and one stamen initiate from a common primordium. Mature stamens curve inwards and cover the ovary in bud. The gynoecium begins as a hemispheric primordium on which two carpellary lobes arise simultaneously. Later in development a single gynocium is formed with two locules and two ovules per locule. Floral morphology suggests a closer affinity with Sapindaceae, although certain features of floral ontogenesis are similar to those observed in certain members of the former Hippocastanaceae, such as Handeliodendron.     

Wrinkled petals and stamens 1, is required for the morphogenesis of petals and stamens in Lotus japonicus

Although much progress has been made in understanding how floral organ identity is determined during the floral development, less is known about how floral organ is elaborated in the late floral developmental stages. Here we describe a novel floral mutant, wrinkled petals and stamens1 （wps1）, which shows defects in the development of petals and stamens. Genetic analysis indicates that wpsl mutant is corresponding to a single recessive locus at the long arm of chromosome 3. The early development of floral organs in wpsl mutant is similar to that in wild type, and the malfunction of the mutant commences in late developmental stages, displaying a defect on the appearance of petals and stamens. In the mature flower, petals and stamen filaments in the mutant are wrinkled or folded, and the cellular morphology under L1 layer of petals and stamen filaments is abnormal. It is found that the expression patterns of floral organ identity genes are not affected in wpsl mutants compared with that of wild type, consistent with the unaltered development of all floral organs. Furthermore, the identities of epidermal cells in different type of petals are maintained. The histological analysis shows that in wpsl flowers all petals are irregularly folded, and there are knotted structures in the petals, while the shape and arrangement of inner cells are malformed and unorganized. Based on these results, we propose that Wpsl acts downstream to the class B floral organ identity genes, and functions to modulate the cellular differentiation during the late flower developmental stages.     

三白草(三白草科)花部器官发生

In the present study,floral organogenesis of Saururus chinensis was observed and compared with that of 5. cernuus. The two species share essentially similar patterns of floral initiation and stamen development. Their inflorescence produces "common primordia" in acropetal succession on the flanks of the inflorescence meristem. Each primordium bifurcates transversely to form a floral apex above and a bract primordium below. Six stamens arise in three pairs at the floral apex. The median sagittal pair arises first, the lateral distal pair second, and the lateral proximal pair arises last. On the contrary,the initiation of carpels is quite different from each other. In 5. cernuus, the median sagittal pair arises first, and the lateral pair next. In S. chinensis, however, the lateral pair arises first, and the median sagittal pair second. The present study also made a brief generalization using the data obtained from different fields on the relationship of the two species in the genus Saururus, which are dis     

Anatomy and ontogeny of unisexual flowers in dioecious Woonyoungia septentrionalis （Dandy） Law （Magnoliaceae）

Woonyoungia septentrionalis （Dandy） Law is aceae. The floral morphology and structure of the species a dioecious species with unisexual flowers in Magnoliare conspicuously different from other species and are important to the study of floral phylogeny in this family. The floral anatomy and ontogeny were investigated to evaluate the systematic position of W. septentrionalis, using scanning electron microscopy and light microscopy. All of the floral organs are initiated acropetally and spirally. The carpels are of conduplicated type without the differentiation of stigma and style. The degenerated stamens in the female flowers have the same structures as the normal stamens at the earlier developmental stages, but they do not undergo successive development and eventually degenerate. The male floral apex was observed to have the remnants of carpels in a few investigated samples. As the bisexual flower features could be traced both in the male and female flowers in W. septentrionalis, it suggests that the flower sex in Magnoliaceae tends toward unisexual. As well as the unisexual flowers, the reduced tepals and carpels and concrescence of carpels conform to the specialized tendency in Magnoliaceae, which confirms the derived position of W. septentrionalis in this family. As the initiation pattern of floral parts of W. septentrionalis is very similar to other species in this family, it needs further investigation and especially comparison with species in Kmeria to evaluate the separation of Woonyoungia.     

The Earliest Normal Flower from Liaoning Province,China

The early evolution of angiosperms has been a focus of intensive research for more than a century. The Yixian Formation in western Liaoning yields one of the earliest angiosperm macrofioras. Despite multitudes of angiosperm fossils uncovered, including Archaefructus and Sinocarpus, no bona fide normal flower has been dated to 125 Ma （mega-annum） or older. Here we report Callianthus dilae gen. et sp. nov. from the Yixian Formation （Early Cretaceous） in western Liaoning, China as the earliest normal flower known to date. The flower demonstrates a typical floral organization, including tepals, androecium, and gynoecium. The tepals are spatulate with parallel veins. The stamens have a slender filament, a globular anther, bristles at the anther apex, and in situ round-triangular pollen grains. The gynoecium is composed of two stylate carpels enclosed in a fleshy envelope, and develops into a ＂hip＂ when mature. Since the well-accepted history of angiosperms is not much longer than 125 Ma, Callianthus together with Chaoyangia, Archaefructus and Sinocarpus from the Yixian Formation demonstrate a surprisingly high diversity of angiosperms, implying a history of angiosperms much longer than currently accepted.     

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.     

Morphogenesis of Pistillate Flowers of Cercidiphyllum japonicum （Cercidiphyllaceae）

Floral morphogenesis and the development of Cercidiphyllumjaponicum Sieb. et Zucc. were observed by scanning electron microscopy （SEM）. The results showed that the pistillate inflorescences were congested spikes with the flowers arranged opposite. Great differences between the so-called ＂bract＂ and the vegetative leaf were observed both in morphogenesis and morphology. In morphogenesis, the ＂bract＂ primordium is crescent-shaped, truncated at the apex and not conduplicate, has no stipule primordium at the base but does have some inconspicuous teeth in the margin that are not glandular. The leaf primordium is triangular, cycloidal at the apex, conduplicate, has two stipule primordia at the base, has one gland-tooth at the apex occurring at first and some gland-teeth in the margin that occur later. In morphology, the ＂bract＂ is also different to the vegetative leaf in some characteristics that were also illustrated in the present paper. Based on the hypothesis that the bract is more similar to the vegetative leaf than the tepal, we considered that the so-called ＂bract＂ of C.japonicum might be the tepal of the pistillate flower in morphological nature. Therefore, each pistillate flower contains a tepal and a carpel. We did not find any trace of other floral organs in the morphogenesis of the pistillate flower. Therefore we considered that the unicarpellate status of extant Cercidiphyllum might be to highly reduce and advance characteristics that make the extant Cercidiphyllum isolated from both fossil Cercidiphyllum-like plants and its extant affinities.     

Conservation of the WD-repeat, microtubule-binding protein, EMAP, in sea urchins, humans, and the nematode C. elegans

The echinoderm microtubule-associated protein (EMAP) is the most abundant microtubule-binding protein in the first cleavage mitotic apparatus in sea urchin embryos. The first goal of this study was to determine whether there is sufficient EMAP in the egg and embryo to modify microtubule dynamics during the early cleavages divisions and whether EMAP functions at a specific time or place in the embryo. To accomplish this goal, we examined the relative abundance, tissue distribution, and temporal pattern of EMAP expression during embryonic development. The second goal of this study was to identify important functional domains within the EMAP coding sequence. A conserved sequence might reveal a potential microtubule-binding domain. We cloned, sequenced and compared overlapping EMAP cDNAs from two different sea urchin species that diverged approximately 80 million years ago, and compared these with cDNA sequences from a vertebrate and nematode species. From quantitative immunoblots, we determined the EMAP concentration in eggs to be 4 μM. The steady-state levels of EMAP mRNA and protein accumulated during development, and all three germ layers expressed EMAP. During the early stages of development, EMAP and tubulin were both abundant in the ectoderm, mesoderm and endoderm. However, during late gastrulation and the formation of the early pluteus larvae, EMAP was enriched in the mesoderm, while tubulin staining was most abundant in the archenteron. These results indicate that EMAP may have tissue-specific functions in the late stage embryo. To identify conserved functional domains, we compared the predicted amino acid sequence encoded by Strongylocentrotus purpuratus and Lytechinus variegatus EMAP cDNAs, and determined that these two sea urchin EMAPs were 95% conserved and shared an identical domain organization. A parsimonious analysis of these sea urchin protein sequences, as well as human and C. elegans EMAP sequences was used to construct a gene tree. Together these results suggest that EMAP is an important microtubule protein required at all developmental stages of sea urchins, and whose cellular function may be conserved amongst metazoans. Received: 2 March 1999 / Accepted: 28 June 1999     

An Arabidopsis embryonic lethal mutant with reduced expression of alanyl—t RNA synthetase gene

In present paper,one of the T-DNA insertional embryonic lethal mutant of Arabidopsis is identified and designated as acd mutant.The embryo developmant of this mutant is arrested in globular stage,The cell division pattern is abnormal during early embryogenesis and results in distubed cellular differentiation.Most of mutant embryos are finally degenerated and aborted in globular stage,However,a few of them still can germinate in agar palte and produce seedlings with shoter hypoctyl and distorted shoot meristem.To understand the molecular basis of the phenotype of this mutant,the joint fragment of T-DNA/plant DNA is isolated by plasmid rescue and Dig-labeled as probe for cDNA library screening.According to the sequence analysis and similarity searching,a 936 bp cDNA sequence(EMBL accession #:Y12555)from selectoed positive clone shows a 99.8%(923/925bp) sequence homolgy with Alanyl-tRNA Synthetase(AlaRS) gene of Arabidopsis thaliana.Furthermore,the data of in situ hybridization experiment indicate that the expression of Ala RS gene is weak in early embryogenesis and declines along with globular embryodevelopment in this mutant Accordingly,the reduced expression of Ala RS gene may be closely related to the morphological changes in early embryogenesis of this lethal mutant.     

Secretory Cavity Development and Its Relationship with the Accumulation of Essential Oil in Fruits of Citrus medica L. var. sarcodactylis （Noot.） Swingle

The developmental types of secretory cavities in Citrus remain controversial. The relationship between secretory cavity development and the accumulation of essential oil in fruits of Citrus species is also unknown. In order to develop better insights into these problems, histological, histochemical, and cytochemical methods were used to investigate secretory cavity development and the accumulation of essential oll at different developmental stages of fruits of Citrus medica L. var. sarcodactylis （Noot.） Swingle. The results indicate that the secretory cavity of the variety seemed to originate from an epidermal cell and a subepldermal cell. These two cells underwent successive divisions, resulting In the formation of two parts： （Ⅰ） a conical cap; and （Ⅱ） a globular gland. The formation of the lumen was schlzolysigenous. Regular changes in the size of vacuoles and the accumulation of essential oil were revealed during the process of secretory cavity development. In addition, when fruits were a light yellow or golden color, the structure of secretory cavities was well developed and the content of essential oil in a single fruit reached a maximum. It would be most appropriate to collect the fruit as a medicinal material at this time.     

Floral Morphogenesis of Anemone rivularis Buch.-Ham. ex DC. var.flore-minore Maxim. (Ranunculaceae) with Special Emphasis on Androecium Developmental Sequence

The floral morphogenesis and androecium developmental sequence of Anemone rivularis Buch.-Ham. ex DC. var. flore-minore Maxim. were observed under a scanning electron microscope (SEM)and by means of histological methods in order to expand our knowledge of the morphogenesis and development of the floral organs of the Ranunculaceae. The initiation of the floral elements is a centripetal spiral and the direction of the spiral is clockwise or anti-clockwise. However, the development of the androecium is highly unusual: in a longitudinal series of four stamens, the second stamen develops first from the inner to outer, then the third one, the fourth one and the first one in turn. The microsporogenesis and anther maturation follows the same developmental sequence. The tepals are different from the bracts and the stamens in both shape and size in the early developmental stage, but there is no difference between the stamens and carpels in the early developmental stage. Therefore, we established a spatio-temporal process of the floral morphogenesis of4. rivularis var.flore-minore and offer another meaning of the floral diversity patterns attributed to the level of the genus.     

Early floral development of Pentaphylacaceae (Ericales) and its systematic implications

Early floral development of four species from the genera Anneslea, Cleyera, Eurya, and Ternstroemia of Pentaphylacaceae, was studied comparatively using scanning electron microscopy. Together with earlier studies in Euryodendron and Adinandra, 6 out of 12 genera of Pentaphylacaceae have now been studied for their floral development. The usually pentamerous flowers of these taxa share a number of developmental features: the perianth organs appear in a clockwise or anticlockwise spiral sequence on the floral apex with relatively long plastochrons between successive organs, resulting in conspicuous size differences among perianth organs during early developmental stages. The early development of the usually polystemonous androecium is characterized by an indistinct ring-primordium and a mostly concave floral apex; individual stamens appear subsequently on this ring-primordium. However, further development of the androecium differs conspicuously among taxa and we describe three main developmental patterns for the family including features such as centripetal stamen whorls and stamens fascicles. Unusual features of floral development and organization of Pentaphylacaceae include: (1) a pronounced spiral sequence of organ appearance during early floral development in perianth and androecium; (2) the occurrence of paired organs in the corolla and the androecium of some species; (3) sepals and petals that are positioned opposite from each other in the genera Anneslea and Ternstroemia; and (4) a concave floral apex at the beginning of androecium development. From a systematic point of view our results clearly support a close relationship between Anneslea and Ternstroemia and also suggest a closer relationship among Adinandra, Cleyera, and Euryodendron on the one hand and between Eurya and Visnea on the other. Further, our developmental study stresses the differences between Pentaphylacaceae and Theaceae, which earlier where thought to form a natural group of plants. While high stamen numbers are achieved via centripetal pattern of stamen formation in the former family, stamens are formed centrifugally in the latter.     

黄连属(毛茛科)花的形态发生

本文运用扫锚电子显微镜(SEM)观察了黄连属(Coptis)植物花的形态发生和发育过程, 结果表明, 该属植物所有的花部器官均为螺旋状发生, 雄蕊为向心式发育, 花瓣原基有微弱的延迟发育, 心皮原基为对折型(即马蹄形), 子房为半封闭类型, 子房柄是在发育过程中形成的。通过与其它具T-型染色体类群在花形态发生上的比较, 认为黄连属表现出了某些原始的性状, 这一结果与分子系统学研究认为黄连属为毛茛科的基部类群的结论一致。     

Floral Developmental Evidence for the Systematic Relationships of Tropaeolum(Tropaeolaceae)

The floral ontogeny of three species of Tropaeolum was studiedusing scanning electron microscopy to find morphological evidencefor discussing the systematic position of the family. The initiationof the androecium is highly unusual: there are always eightstamens which arise (1) either in a spiral sequence startingwith the stamen opposite sepal four, running in a directionopposite to the sequence of the sepals, and with reversals inthe direction of the spiral, or (2) as a sequence of pairedand unpaired stamens. The floral symmetry changes twice duringthe development of the flower, from polysymmetrical at sepaland petal initiation, through oblique monosymmetry at stameninitiation, and ending with median monosymmetry in later developmentalstages. The occurrence of median monosymmetry is a late-developmentalevent and is caused by the initiation of a hypanthial spur,and the unequal growth of the petals and styles. The originfor the unusual sequence of stamen initiation reflects a trendaffecting the whole flower which is linked with the changingpatterns of floral symmetry. Octandry is enhanced by multiplecauses, such as the loss of two stamens in an originally diplostemonousandroecium and the regulating pressure of the gynoecium. Thechange in symmetry during ontogeny is significant for discussingthe systematic position of Tropaeolaceae in comparison withthe glucosinolate-producing taxa and the Sapindales. The combinationof an androecium with eight stamens and oblique monosymmetryis either a single event in evolution and links Tropaeolum withthe Sapindales, or it has evolved at least twice, once in theSapindales, and once in a clade comprising Tropaeolaceae, Akaniaceaeand Bretschneideraceae. Morphological data support a sistergroup relationship of the three latter families, which is inline with macromolecular studies. Copyright 2001 Annals of BotanyCompany Tropaeolum, Tropaeolaceae, Glucosinolate clade, Sapindales, oblique monosymmetry, androecium, octandry, floral development, phylogeny     

Early floral development and androecium organization in Fouquieriaceae (Ericales)

Early floral development with focus on the androecium was studied with the help of scanning electron microscopy and serial microtome sectioning in Fouquieria columnaris and F. splendens. Perianth organs appear in a spiral pattern on the floral apex. The spiral may be a clockwise or anti-clockwise. The androecium is best interpreted as two-whorled with all the stamens arranged in a single series. In F. splendens, two or more of the five epipetalous stamen positions are doubled, i.e. they are occupied by stamen pairs. Unusual features in the floral development of Fouquieriaceae include (1) a strong spiral component even in whorled organ categories and (2) a pronouncedly asymmetric floral apex during an early phase of floral development. From a phylogenetic point of view, it seems plausible that the common ancestor of Fouquieriaceae and its sister family Polemoniaceae was characterized by two alternating, pentamerous stamen-whorls.     

Floral development of Bougainvillea spectabilis Willd., Boerhaavia diffusa L. and Mirabilis jalapa L. (Nyctaginaceae)

Three morphological problems were investigated in three species of the Nyctaginaceae: epiphylly, phyllotaxis and placentation. Epiphylly, which occurs in Bougainvillea spectabilis , is the result of ontogenetic displacement resulting from the activity of an intercalary meristem at the base of the floral bract and the floral bud. Floral development of Bougainvillea spectabilis was compared with that of Boerhaavia diffusa and Mirabilis jalapa . Considerable variation occurs with regard to the number and arrangement of stamens. Five stamens are initiated simultaneously, alternate to the petals, in Mirabilis . In Bougainvillea , eight stamens arise sequentially at divergence angles suggestive of a 3/8 spiral. No developmental evidence was found to support the derivation of the eight stamens from a two whorled pentamerous androecium. Boerhaavia normally has only two stamens which most frequently are initiated toward opposite sides of the floral apex, but may also be formed in a 2/5 to 3/8 divergence. In some flowers only one or three stamens are formed. The gynoecium is formed in the same way in all three species: growth occurs in a crescent-shaped zone at the periphery of the floral apex thus producing the gynoecial wall. The single ovule, which is basal in the mature gynoecium, is formed from the gradual upgrowth and transformation of the floral apex and is developmentally terminal. Even the two-layered tunica is maintained as the floral apex is transformed into the ovule primordium. If 'carpel' is defined traditionally as a folded megasporophyll which bears and encloses ovule(s) then carpels are not present in the gynoecia of the three species studied. If 'carpel' is re-defined as an appendage which encloses ovule(s), then the gynoecia of the Nyctaginaceae are carpellate. However, the ovules remain cauline regardless of which definition is adopted.     

Floral morphogenesis of Wikstroemia delavayi (Thymelaeaceae) and its phylogenetic implication

Floral morphogenesis of Wikstroemia delavayi Lecomte was investigated by scanning electron microscope (SEM) and compared with its allied groups. Initiation and early development of floral parts in W. delavayi followed unidirectional sequences from the abaxial side to the adaxial side. Because the floral parts grew faster at the adaxial side than at the abaxial one in following development, the zygomorphic pattern in the early development changed and finally became an almost actinomorphic form at anthesis. The disc was initiated from the abaxial base of the floral tube and itslobes were alternate with lower whorl stamens. According to this initiatial and developmental pattern, it is reasonable to interpret the disc as a part of the androecium rather than a modification of the receptacle. The located position and development of the disc was correlative with the development of other floral organs, which might provide insight to delimit Wikstroemia and Daphne based on different floral developmental pattern that might exist between the two genera. The developmental process of W. delavayi indicated that the syncarpous and uniloculate gynoecium was in fact bicarpellate, which consisted of a fertile carpel and a sterile one. It was pseudomonomerous. Even though the ovary in both Wikstroemia and Daphne was uniloculate, the location of the ventral bundles in the ovary was obviously different from each other according to data to date. In this respect, further investigation is undertaken between the two genera.     

澜沧荛花(瑞香科)的花部形态发生 及其系统学意义

通过扫描电镜对澜沧荛花Wikstroemia delavayi花部的形态发生过程进行了观察和分析,旨在为该属的系统学研究提供花部发育形态学资料。澜沧尧花花部的发生和早期发育呈远轴面向近轴面的顺序,但这一式样由于近轴面的器官在早期发育之后生长加速发生了转变。因此,花开放时所表现的所谓辐射对称,显然是由同一轮器官的异率生长所导致的次生现象。花盘发生于花萼筒基部的远轴面上,与花萼、雄蕊的发生间隔时间较长。花盘原基在下轮雄蕊着生处凹陷或间断,与之相对应,花盘裂片与下轮雄蕊呈互生。由此,花盘显然不是花托的一部分,也不是象花萼、雄蕊和心皮一样的独立结构,将其解释为雄蕊群的一部分更合理。花盘的发生和早期发育及其着生位置同其他花部器官的发生和发育式样具有明显的相关性,这种相关性对进一步阐明瑞香属Daphne和荛花属Wikstroemia的系统发育关系具有—定意义。根据对雌蕊群的发生和发育过程观察,该种的子房是由一个近轴面的可育心皮和一个远轴面的不育心皮融合而成的单室子房,为假单心皮雌蕊。尽管荛花属和瑞香属均属于单室产房,但澜沧荛花的子房维管束中的腹束排列于中轴位置,而目前资料显示瑞香属植物的腹束接近于侧膜位置,这方面仍需进一步研究。     

Floral development of Kingdonia (Ranunculaceae s. l., Ranunculales)

The flower of Kingdonia has a terminal position, thus the rhizome is sympodial. The floral organs initiate in spiral phyllotaxis. The androecium is centripetal in initiation but the sterile stamens are retarded in development compared with the fertile ones. The apex of the young carpel does not participate in the conduplication. The floral organs have single vascular traces and unilacunar nodes.The study was supported by the National Nature Science Foundation of China (No. 30370095 and 30130030).     

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.