LABILE SEX EXPRESSION IN ANDROMONOECIOUS SOLANUM HIRTUM: FLORAL DEVELOPMENT AND SEX DETERMINATION

Sex expression (the proportions of hermaphrodite and staminate flowers produced) of the andromonoecious species Solatium hirtum is labile, and this lability of whole plant sex expression is due to labile sex expression of individual floral buds. In this paper I examine the developmental processes that underlie the differences in floral sex expression of hermaphrodite and staminate flowers of Solarium hirtum, focusing particularly on the processes responsible for the observed lability of floral sex expression. Differences in bud growth rate and relative growth of floral organs in these buds are evident at about the time of megasporocyte meiosis (11–12 days before anthesis). However, gynoecial sterility in staminate buds does not occur until just 6–7 days before anthesis. At this time, abnormalities in ovule development occur in staminate buds: the ovules begin to appear necrotic, the integumentary tapetum collapses, and the megagametophytes of many ovules cease normal development. These observations are consistent with the predictions of labile floral development.     

Fruit-Set Induced Changes in the Sex of Flowers in Caesalpinia calycina (Leguminosae)

Abstract: The proportions of hermaphrodite to staminate flowers in inflorescences of the andromonoecious species Caesalpinb calycina vary. Analysis of fruit position along the flowering rhachis, and also of the sex of floral buds in inflorescences with fruits set in different positions, indicate that fruiting success influences the sex of flowers in distal positions along the rhachis. Other reports of andromonoecy in caesalpinoid species are examined with reference to floral sex lability and the influence of fruit set.     

Diversification of andromonoecy in Solanum section Lasiocarpa (Solanaceae): the roles of phenotypic plasticity and architecture

Quantitative analyses of sexual expression show extensive interspecific variation in the strength of andromonoecy (proportions of hermaphroditic and staminate flowers) among Solanum species in the monophyletic section Lasiocarpa. The roles of phenotypic plasticity and inter- and intra-inflorescence architecture in the diversification of andromonoecy within this small clade were analyzed. Four species that represent a range of expression of andromonoecy were examined. Staminate flowers produced within inflorescences ranged from 3% (S. candidum) to 7% (S. ferox) in weakly andromonoecious species and from 39% (S. pseudolulo) to 60% (S. quitoense) in more strongly andromonoecious species. Manipulation of fruit set on clonal replicates of multiple genotypes demonstrated variation among species for phenotypic plasticity. The strongly andromonoecious species, S. pseudolulo and S. quitoense, were not plastic and produced a large proportion of staminate flowers regardless of fruiting treatment, whereas S. candidum and S. ferox were phenotypically plastic and produced significantly more staminate flowers in the presence of developing fruit. Staminate flower production of all four species varied both within and among inflorescences. A greater proportion of staminate flowers were produced in distal (later produced) inflorescences. Within inflorescences, hermaphroditic flowers occurred in basal positions, whereas staminate flowers, when produced, occurred more distally. This pattern of staminate flower production is qualitatively the same in all species investigated; however, quantitative variation in the transition from hermaphroditic to staminate flower production within and among inflorescences is associated with variation in the strength of andromonoecy. At least three factors have contributed to the diversification of andromonoecy in section Lasiocarpa including the presence or absence of phenotypic plasticity in response to fruit set, quantitative variation in intra- and inter-inflorescence architectural effects, and total flower production.     

DEVELOPMENTAL PLASTICITY,GENETIC VARIATION,AND THE EVOLUTION OF ANDROMONOECY IN SOLANUM HIRTUM (SOLANACEAE)

Studies of andromonoecious species have shown that sex expression (proportions of hermaphrodite and staminate flowers) is quite variable. It is not known, however, whether this variation is due to variation among individuals for genetically fixed patterns of allocation to staminate and hermaphrodite flowers (population level variation) and/or to developmental plasticity of individuals in a heterogeneous environment (organismal level variation). Distinguishing between these two levels of variation is important for understanding the evolution of andromonoecy. This study investigates levels of variation in sex expression in the andromonoecious Solanum hirlum. Sex expression in this species is shown to be plastic among individuals of the same genotype (organismal level variation) and determined, in part, by the resource status of the individual. Among the genotypes examined there is also genetic variation for developmental plasticity. Thus, developmental plasticity can potentially respond to selection, and the evolution of this developmental system may have been instrumental in the establishment and maintenance of andromonoecy in S. hirtum.     

臭椿花器官分化的初步研究

利用扫描电镜(SEM)和光镜(LM)对臭椿花序及花器官的分化和发育进行了初步研究,表明:1)臭椿花器官分化于当年的4月初,为圆锥花序;2)分化顺序为花萼原基、花冠原基、雄蕊原基和雌蕊原基。5个萼片原基的发生不同步,并且呈螺旋状发生;5个花瓣原基几乎同步发生且其生长要比雄蕊原基缓慢;雄蕊10枚,两轮排列,每轮5个原基的分化基本是同步的;雌蕊5,其分化速度较快;3)在两性花植株中,5个心皮顶端粘合形成柱头和花柱,而在雄株中,5个心皮退化,只有雄蕊原基分化出花药和花丝。本研究着重观察了臭椿中雄花及两性花发育的过程中两性花向单性花的转变。结果表明,臭椿两性花及单性花的形成在花器官的各原基上是一致的(尽管时间上有差异),雌雄蕊原基同时出现在每一个花器官分化过程中,但是,可育性结构部分的形成取决于其原基是否分化成所应有的结构:雄蕊原基分化形成花药与花丝,雌蕊原基分化形成花柱、柱头和子房。臭椿单性花的形成是由于两性花中雌蕊原基的退化所造成,其机理有待于进一步研究。     

Sex expression in fruiting male vines of kiwifruit

Summary In three canes on each of five fruiting male kiwifruit vines (A. deliciosa var. deliciosa (A. Chev) Liang and Ferguson), staminate and bisexual flowers were interspersed, so that staminate, bisexual and mixed inflorescences were found. Further variation in inflorescence composition resulted from the abortion of flower buds prior to anthesis. These two sources of variation, in combination, resulted in 19 inflorescence patterns, which varied substantially in frequency among and within vines. Some tendencies were determined. In particular, most of the inflorescences had retained their terminal flower, whereas just over half had retained one or both primary lateral flowers, and in two-thirds of the inflorescences retaining a terminal flower, this was bisexual. The five vines ranged in bisexual flower frequency from 40% to 70%. Phenotypic Gender (P) estimates are given for each vine, based on the frequencies of staminate and bisexual flowers from three canes per vine. Flower development, lability in sex expression and the use of P estimates as part of a selection index in breeding for hermaphrodite kiwifruit cultivars are discussed.     

Is LEAFY a useful marker gene for the flower-inflorescence boundary in the Euphorbia cyathium?

The flower-like reproductive structure of Euphorbia s.l. (Euphorbiaceae) is widely believed to have evolved from an inflorescence, and is therefore interpreted as a special type of pseudanthium, termed a cyathium. However, fuzzy morphological boundaries between the inflorescence, individual flowers, and organs have fuelled the suggestion that the cyathium does not merely superficially resemble a flower but could actually share developmental genetic pathways with a conventional flower. To test this hypothesis, immunolocalizations of FLORICAULA/LEAFY (LFY), a protein associated with floral identity in many angiosperm species, were performed in developing cyathia of different species of Euphorbia. Expression of the LFY protein was found not only in individual floral primordia (as predicted from results in the model organisms Arabidopsis and Anthirrhinum), but also in the cyathium primordium and in the primordia of partial male inflorescences. These results provide further evidence that the evolution of floral traits in pseudanthial inflorescences often involves expression of floral development genes in the inflorescence apex. This finding blurs the conventional rigid distinction between flowers and inflorescences.     

Flower structure and potential bisexuality in Freycinetia reineckei (Pandanaceae), a species of the Samoa Islands

In Freycinetia reineckei the staminate flower (on the staminate spikes) comprises 3 or 4 (sometimes 2) stamens and a pistillode with 2 (sometimes 4) carpellodes, and the pistillate flower (on the pistillate spikes) is formed of a pistil with 2 (sometimes 4) carpels and of 3 or 4 (sometimes 2) staminodes. This perfect floral homology, also observed in all the other species that were studied with both pistillate and staminate material, strongly suggests that the flower of Freycinetia is basically and potentially bisexual, and may explain the occasional sexual lability and bisexuality of that flower (occurrence of both pistillate and staminate inflorescences, and/or of bisexual inflorescences with bisexual flowers and/or unisexual flowers, on the same individuals) in some species, and also the frequent occurrence of bisexual spikes in this species. These may be partitioned into pistillate, staminate, mixed and sterile zones. In the pistillate zones the flowers have the same aspect and structure as the pistillate flowers. In the staminate zones the flowers generally comprise 3 or 4 (sometimes 2) stamens and a ‘semi-pistil’ some have both stamens and staminodes. The semi-pistils are intermediate between pistils and pistillodes in length, aspect and structure, but always have placentas and ovules. In the mixed zones the flowers are generally formed of a pistil and 3 or 4 (sometimes 2) stamens, and are therefore true hermaphrodite flowers; some have both stamens and staminodes. In the sterile zones the flowers comprise a semi-pistil and 3 or 4 (sometimes 2) staminodes. The staminodes are anatomically very similar to the stamens, especially in the staminate, mixed, and sterile zones, in which they exhibit a wide range of variation in length, aspect and structure. The perfect floral homology as generic character on one hand, and the occasional bisexuality both with and without bisexual flowers and other aspects of sex expression (e.g. occurrence of both pistillate and staminate shoots on the same individuals) in some species on the other hand, seem to indicate that Freycinetia is a basically monoecious, sex changing genus.     

Morphogenetic lability of reproductive structures in Ruppia maritima (Ruppiaceae, Alismatales): from two lateral flowers to a terminal flower

Flowers of Ruppia are normally arranged into an open two-flowered spike, but sometimes the two lateral flowers are congenitally united with each other and form a terminal flower-like structure. This developmental abnormality resembles those described in well-investigated mutants of model organisms of developmental genetics such as Arabidopsis Antirrhinum. A study of Ruppia allows investigating morphogenetic lability of this feature in natural populations. These data will be important for understanding evolutionary transitions between open and closed inflorescences. This paper presents first data on frequencies ofterminal flower-like structures in natural populations of Ruppia maritima and first observations of their development. Vascular supply of inflorescences with free and united flowers is compared for the first time. Strong differences in frequencies of occurrence of terminal flower-like structures among examined natural populations are revealed. Data on variation of organ numbers in flowers of plants from different populations allow hypothesizing that increased size of floral primordia is a factor that plays a role in their amalgamation into ajoint primordium of a terminal structure. Vascular system of inflorescences of R. maritima with united flowers is quite similar to the vascular system of a flower and nothing contradicts a hypothesis on terminal position ofthis structure. Transversally inserted stamens in inflorescences with united flowers are usually of inverted polarity. This appears to be the first documented example of an inversion of relative polarity of stamens and carpels in angiosperms.     

Floral ontogeny in Scirpus, Eriophorum and Dulichium (Cyperaceae), with special reference to the perianth

BACKGROUND AND AIMS: Based on molecular phylogenetic analysis, it has been suggested recently that the Cyperaceae comprises only two subfamilies: the Mapanioideae and the Cyperoideae. In most flowers of the Cyperoideae, the whorl of inner stamens is reduced, resulting in tetracyclic flowers. In the more primitive (scirpoid) genera within the Cyperoideae, the perianth consists of two polysymmetric whorls, whereas the perianth parts in the more derived genera have been subject to modifications and/or reduction. Comparative studies of the many silky hairs of Eriophorum and of the eight bristles of Dulichium have given rise to much discussion about their homology. METHODS: The spikelet and floral ontogeny in freshly collected inflorescences was investigated using scanning electron microscopy. KEY RESULTS: Complete floral ontogenies are presented for Scirpus sylvaticus L., Eriophorum latifolium Hoppe and Dulichium arundinaceum (L.) Britton, with special reference to the perianth. The results in S. sylvaticus confirm the trimerous monocot-like organization of the flower. It is used as a model for floral development in Cyperoideae. In the early developmental stages, the androecium of E. latifolium is surrounded by a massive perigonial primordium, from which the many hair-like bristles originate. Consequently, the stamens develop among the hair primordia, more or less simultaneously. The hairs are arranged in whorls, which develop centripetally. The development of the perianth in D. arundinaceum starts with the formation of three initial perianth primordia opposite the stamens. Subsequently, two more abaxial bristle primordia, alternating with the stamens, originate simultaneously with the appearance of three adaxial bristle primordia in the zone where an adaxial inner perianth primordium is expected. CONCLUSIONS: The floral development in E. latifolium and D. arundinaceum can be considered as variations upon the scirpoid floral ontogenetic theme.     

雄全同株植物簇花芹花期性别分配与开花式样

簇花芹(Soranthus meyeri)是古尔班通古特沙漠中常见的、具雄全同株性系统的伞形科多年生早春短命植物。该文对簇花芹花期性比(两性花数/总花数)与植株大小的关系及其开花式样进行了研究, 重点对花期大小依赖的性别资源分配进行了讨论。结果表明: 2006-2008年簇花芹群体水平的性比分别为0.69 ± 0.03、0.62 ± 0.03和0.69 ± 0.02, 彼此间无显著差异( p > 0.05), 表明其性比是相对稳定的, 可能受遗传因素的控制。雄花生物量与花粉量均比两性花的小, 说明产生雄花比产生两性花所需资源少。一级复伞形花序比二级复伞形花序具有较多的两性花, 说明前者易从植株上获得资源用于增加雌性适合度; 而后者产生较多的雄花以避免在雌性功能上资源投入的浪费, 增大花展示以吸引更多传粉者来增加花粉输出总量, 提高其整体适合度。植株水平的性比与地上营养器官的生物量间呈正相关关系, 说明较大个体对雌性功能的投资较大, 雌性繁殖成功受资源限制。复伞形花序内各伞形花序几乎同时向心开放, 且所有两性花及花序均为雄性先熟, 雌雄阶段完全分离, 但一级复伞形花序比二级复伞形花序早开放约5天, 彼此开花重叠期约为1天。这些特征对于一级复伞形花序进行异株异花授粉以及植株内不同级别花序间的同株异花传粉、避免雌雄功能间的干扰具有重要意义。     

The role of growth regulators in the differentiation of flowers and inflorescences

Growth regulators participate in the differentiation of floral parts, determining the developmental path of the respective type of inflorescence. The effect depends on the expression of the peculiarities of floral part differentiation, the recognition of the character of endogenous substances in certain stages and the choice of the suitable regulator for application. In the primitive flower ofPapaver petals and stamens are formed from the peripheral meristem with a lower content of auxins and a higher level of gibberellic substances. The pistil arises later from central tissues with a higher level of auxins and inhibitory substances. The stamens are more sensitive to the higher level of auxin substances, and by a suitable application of GA₃ and BAP they can be transformed into petals; in this way double flower forms arise. In the differentiation of floral parts ofCampanula, Rosa andMelandrium similar regularities assert themselves in time successions, but in another spatial arrangement. Sex differentiation of diclinous flowers ofMelandrium is based on differences in heterochromosomes XY and XX. The rise of the zygomorphic flower ofVeronica is accompanied by a different distribution of endogenous substances which affect the development of petals, stamens and the pistil. The differentiation of flowers in the racemose inflorescence occurs in the acropetal succession, and lateral primordia inCampanula develop into actinomorphic regular flowers, whereas inDigitalis they are zygomorphic and only the terminal flower is peloric. In the initial phases the staminate tassel and the pistillate ear in maize are identical. Earlier differentiation of the terminal pistillate tassel is connected with a higher level of gibberellins and the later development of the lateral pistillate ear is accompanied by the increase in auxin-like substances and inhibitions. Similar correlations were found in the development of staminate catkins and the differentiation of pistillate flowers in terminal buds ofJuglans regia. By the application of auxin-like substances it is possible to achieve the transformation of primordia of the staminate tassel into the pistillate ear in maize or to regulate the number of staminate catkins and pistillate flowers on twigs of the walnut tree. In the capitulum of the sunflower differences arise between peripheral pistillate ray flowers and hermaphrodite tubular ones. By applying GA₃ and BAP the number of ray flowers is increased. If the normal course of inflorescence differentiation is affected with a suitable type of regulator, a range of floral abnormalities appears which permit to assess the intervention in different developmental stages and the reaction of the primordium to the applied type of regulator. Abnormalities also suggest some phylogenetic correlations.     

FLORAL DEVELOPMENT IN MYRISTICA (MYRISTICACEAE)

Myristica fragrans and M. malabarica are dioecious. Both staminate and pistillate plants produce axillary flowering structures. Each pistillate flower is solitary, borne terminally on a short, second-order shoot that bears a pair of ephemeral bracts. Each staminate inflorescence similarly produces a terminal flower and, usually, a third-order, racemose axis in the axil of each pair of bracts. Each flower on these indeterminate axes is in the axil of a bract. On the abaxial side immediately below the perianth, each flower has a bracteole, which is produced by the floral apex. Three tepal primordia are initiated on the margins of the floral apex in an acyclic pattern. Subsequent intercalary growth produces a perianth tube. Alternate with the tepals, three anther primordia arise on the margins of a broadened floral apex in an acyclic or helical pattern. Usually two more anther primordia arise adjacent to each of the first three primordia, producing a total of nine primordia. At this stage the floral apex begins to lose its meristematic appearance, but the residuum persists. Intercalary growth below the floral apex produces a columnar receptacle. The anther primordia remain adnate to the receptacle and grow longitudinally as the receptacle elongates. Each primordium develops into an anther with two pairs of septate, elongate microsporangia. In pistillate flowers, a carpel primordium encircles the floral apex eventually producing an ascidiate carpel with a cleft on the oblique apex and upper adaxial wall. The floral ontogeny supports the morphological interpretation of myristicaceous flowers as trimerous with either four-sporangiate anthers or monocarpellate pistils.     

Morphogenetic lability of the <Emphasis Type="Italic">Ruppia maritima</Emphasis> (Ruppiaceae,Alismatales) reproductive organs: From two lateral flowers to a terminal flower

Flowers of Ruppia are usually arranged into an open two-flowered spike, but sometimes two lateral flowers are congenitally united with each other forming a terminal flower-like structure. This deviation from the morphogenesis of reproductive structures typical of Ruppia resembles those described in well-studied mutants of the model organisms of developmental genetics, such as Arabidopsis and Antirrhinum. A study of Ruppia allows the morphogenetic lability of this trait to be traced in natural populations. These data are important for understanding the evolutionary transition from open to closed inflorescences. This paper describes the first data on the frequencies of terminal flower-like structures in natural populations of Ruppia maritima as well as on the specific features of their morphogenesis. The vascular supply in the inflorescences with free and fused flowers is also compared for the first time. It has been demonstrated that the frequency of inflorescences with fused flowers considerably varies between different populations. The data on variation in the number of organs in flowers of plants from different populations suggest that an increased size of floral primordia is a factor enhancing their fusion into a joint primordium of the terminal structure. The vascular system of the R. maritima inflorescences with united flowers is similar to the vascular system of a single flower; moreover, nothing contradicts the hypothesis on a terminal position of this structure. The R. maritima inflorescences with united flowers frequently contain transversal stamens with an inverted polarity. Presumably, this is the first case of recorded inversion of relative polarity of stamens and carpels in angiosperms.     

Pistillate and staminate flower development in dioecious Pistacia vera (Anacardiaceae)

The development of staminate and pistillate flowers in the dioecious tree species Pistacia vera L. (Anacardiaceae) was studied by scanning electron microscopy with the objective of determining organogenetic patterns and phenology of floral differentiation. Flower primordia are initiated similarly in trees of both sexes. Stamen and carpel primordia are initiated in both male and female flowers, and the phenology of organ initiation is essentially identical for flowers of both sexes. Vestigial stamen primordia arise at the flanks of pistillate flower apices at the same time functional stamens are initiated in the staminate flowers. Similarly, a vestigial carpel is initiated in staminate flowers at the same time the primary, functional carpel is initiated in pistillate flower primordia. Differences between the two sexes become apparent early in development as, in both cases, development of organs of the opposite sex becomes arrested at the primordial stage. Male flowers produce between four and six mature functional stamens and female flowers produce a gynoecium with one functional and two sterile carpels.     

Sexual dimorphism and gynoecium size variation in the andromonoecious shrub Caesalpinia gilliesii

The degree of sexual dimorphism in flowers and inflorescences can be evaluated early in flower development through the study of floral organ size co-variation. In the present work, the gynoecium-androecium size relationship was studied to assess the degree of sexual expression in flowers and inflorescences of the andromonoecious shrub Caesalpinia gilliesii. The co-variation pattern of floral organ sizes was compared between small and large inflorescences, under the hypothesis that inflorescence size reflected differential resource availability. Also, staminate and perfect flowers were collected from three populations and compared on the basis of gynoecium, ovule length, filament length, pollen size and number. The obtained results indicated that staminate and perfect flowers differed only in the gynoecium and ovule length, whereas filament length, pollen size, and number varied across populations. The gynoecium size was smaller and its variability was much higher in staminate than in perfect flowers, as explained by a recent hypothesis about pollinator-mediated gynoecium size selection acting upon perfect flowers. The analysis of the gynoecium-androecium size relationship during flower development, revealed a dissociation of gynoecium growth relative to other floral structures in some buds. Lower gynoecium-androecium regression slopes and smaller gynoecia length characterized smaller inflorescences, thus reflecting the fact that sexual expression was more male-biased. This trend is in agreement with a differential resource-related response at the inflorescence level, however, post-mating resource allocation and the inclusion of other modular levels may also help us to understand the variation in sexual dimorphism in this species.     

Unisexual flowers as a robust synapomorphy in Cariceae (Cyperaceae)? Evidence for bisexual flowers in Schoenoxiphium

Cariceae, the largest tribe within Cyperaceae, comprises about 2000 species in five genera. Cariceae is usually considered to be distinct from other Cyperaceae by the presence of exclusively unisexual flowers and by the arrangement of the pistillate flowers in single-flowered spikelets that are enclosed by the flask-like spikelet prophyll (utricle or perigynium). The nature of several morphological features of the Cariceae inflorescence remains controversial. The staminate reproductive units, as well as earlier reported bisexual reproductive units in Schoenoxiphium have been considered to be reduced partial inflorescences, or flowers. Aims of this study are to test both interpretations, based on a floral ontogenetic investigation. Moreover, for the first time, detailed SEM micrographs are presented of the inflorescence and floral development and of bisexual flowers in Schoenoxiphium. We propose that ‘inhibition of bisexuality’ is a more robust synapomorphy in Cariceae than ‘presence of only unisexual flowers’.     

Flower-like heads from flower-like meristems: pseudanthium development in <Emphasis Type="Italic">Davidia involucrata</Emphasis> (Nyssaceae)

Flower-like inflorescences (pseudanthia) have fascinated botanists for a long time. They are explained as condensed inflorescences implying that the pseudanthium develops from an inflorescence meristem (IM). However, recent developmental studies identified a new form of reproductive meristem, the floral unit meristem (FUM). It differs from IMs by lacking acropetal growth and shares fractionation, expansion and autonomous space filling with flower meristems (FM). The similarity among FUMs and FMs raises the question how far flower-like heads originate from flower-like meristems. In the present paper, pseudanthium development in Davidia involucrata is investigated using scanning electron microscopy. D. involucrata has pincushion-shaped heads composed of densely aggregated, perianthless flowers and associated with two large showy bracts. Early developmental stages show a huge naked FUM. The FMs appear almost simultaneously and lack subtending bracts. With ongoing FUM expansion new space is generated which is immediately used by further FM fractionation. The heads have only staminate flowers or are andromonoecious with staminate and a single perfect flower in oblique position. All FMs lack perianth structures and fractionate a variable number of stamen primordia. The perfect FM is much larger than the staminate FMs and forms a syncarpous gynoecium with inferior ovary. Pseudanthium development in D. involucrata confirms the morphogenetic similarity to FMs as to acropetal growth limitation, meristem expansion and fractionation. It thus should not be interpreted as a condensed inflorescence, but as a flower equivalent. Furthermore as the FUM develops inside a bud, its development is considered to be influenced by mechanical pressure. The oblique position of the perfect flower, the developmental delay of the proximal flowers, and the variable number of stamens which were observed in the pseudanthium development, can be caused by mechanical pressure. Next to the Asteraceae, D. involucrata offers a further example of a pseudanthium originating from a FUM. More knowledge on FUMs is still needed to understand diversification and evolution of flower-like inflorescences.     

HELICAL FLORAL ORGANOGENESIS IN GLEDITSIA,A PRIMITIVE CAESALPINIOID LEGUME

In order to determine the extent of floral ontogenetic differences among species of a genus, six species of Gleditsia were studied. Gledilsia is one of only two leguminous genera known in which there is completely helical succession of floral organs. Floral ontogeny was compared in three species (Gleditsia amorphoides, G. aquatica, and G. triacanthos), and late stages in six species (including the first three plus G. caspica, G. delavayi, and G. japonica). Other unusual primitive developmental features include the unequal-sized flower primordia which produce flowers of variable merosity. Order of floral development is also loosely controlled, so that flowers of different growth stages are intermixed in the inflorescence. Variable features include the occurrence of floral bracts, merosity of flowers, number of organs, and position of the first organ (sepal) initiated. The inflorescence type, while usually a raceme, often has lateral branches near the base, or fascicles of flowers at some points. A terminal flower often is present, although not in all species. Sex of flowers and inflorescences also varies, although floral initiation tends to include both stamens and carpel primordia. Suppression of one or the other may occur at different stages of development. Carpel orientation also varies; the cleft may be tilted or inverted occasionally. It is proposed that absence of subtending floral bracts influences development so as to favor radial symmetry and establishment of other “chaotic” characters seen in Gledilsia flowers.