Flower Development in Silene: Morphology and Sequence of Initiation of Primordia

The initiation and development of the flower of Silene coeli-rosawas followed by examining apices by scanning electron microscopy.The sepals, stamens and carpets are initiated in a spiral sequence,the direction of the spiral king the opposite of the acropetalhelix of unequal axillary buds at the nodes below the flower.The petals are initiated almost simultaneously and at the sametime as the first few stamens. The change in phyllotaxis fromopposite and decussate in the vegetative shoot to spiral inthe flower occurs with the displacement of the first two sepalsaway from the mid-line of the apex and towards the axillarybud at the node below the flower. The sizes of the sepals andstamens are a function of their age since initiation but thepetals grow more slowly. The Silene flower can be interpretedas a shoot bearing primordia with associated axillary primordia,some of the latter being precocious in their development. Silent weli-rosa, flower initiation, flower development, phyllotaxis, primordia     

Rates of Growth and Primordial Initiation During Flower Development in Silene at Different Temperatures

The growth of the flower and its constituent parts was measuredin Silene coeli-rosa plants, induced at 13, 20 and 27 °C,in order to try and identify those processes which consistentlyoccurred and would therefore be more likely to be essentialfor flower formation. The increased growth rate of the apical dome just before orabout the time of sepal initiation was not maintained in theflower, the growth rate of which was comparable to that of avegetative apex until all the carpels had been initiated, whenit decreased further. The primordia of the same whorl all hadsimilar growth rates so that the relative sizes of the primordiareflected their relative ages since their initiation. The relativegrowth rate of the stamens was the same (13 and 20 °C) orless (27 °C) than that of the sepals, but the relative growthrate of the petals was lower than either. The growth rate ofthe flower axis was least at the sepal node and increased bothdistally and proximally from this region. The plastochron during sepal initiation was shorter than forleaf initiation and tended to be shorter still during initiationof stamens and petals. Increasing temperature increased therate of primordial initiation but at 27 °C the growth ratesof the primordia were lowest although the rates of primordiainitiation were highest. The form of the flower, as exemplifiedby the relative sizes of the primordia at the moment when allcarpels had been initiated, was constant despite the differinggrowth rates and sizes of the primordia on initiation in differenttemperatures. It is concluded that neither the initiation ofthe primordia in the flower nor the form of the flower is determinedprimarily by the relative growth rates of its component parts. Silene coeli-rosa, flower development, primordia initiation, growth     

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.     

突脉金丝桃的花器官发生及其系统学意义

利用扫描电镜（SEM）观察了突脉金丝桃（Hypericum przewalskii）（金丝桃科）的花部器官发生发育过程。结果表明,突脉金丝桃2枚苞片原基首先发生,花原基在苞片原基的包裹中完成发育。在苞片原基发生后,5枚萼片原基沿2/5圆周依次发生。萼片原基发生近完成时,5枚雄蕊—花瓣共同原基在萼片原基之间的角隅处近同时发生,此后,雄蕊—花瓣共同原基下部向外伸展形成花瓣原基,上部向上凸起形成与花瓣原基相对的雄蕊原基,之后雄蕊原基由内向外依次分化发育产生次生雄蕊原基,随着次生雄蕊原基的发育和数目的增多,形成了5束雄蕊。次生雄蕊原基发生的同时,5枚心皮原基近同时发生。突脉金丝桃雄蕊束的发生方式表明,金丝桃属的雄蕊束可能起源于5基数的单轮雄蕊。金丝桃科与藤黄科植物花瓣及雄蕊原基发生方式的显著不同,支持了APG Ⅲ系统将金丝桃亚科从藤黄科中独立为金丝桃科的观点。     

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.     

NORMAL FLORAL ONTOGENY AND COOL TEMPERATURE-INDUCED ABERRANT FLORAL DEVELOPMENT IN GLYCINE MAX (FABACEAE)

Floral onset in soybean (Glycine max cv. Ransom) is characterized by precocious initiation of axillary meristems in the axils of the most recently initiated leaf primordium. During floral transition, leaf morphology changes from trifoliolate leaf with stipules, to a three-lobed bract, to an unlobed bract. Soybean flowers initiated at 26/22 C day/night temperatures are normal, papilionaceous, and pentamerous. Sepal, petal, and stamen whorls are initiated unidirectionally from the abaxial to adaxial side of the floral apex. The median sepal is located abaxially and the median petal adaxially on the meristem. The organogeny of ‘Ransom’ flowers was found to be: sepals, petals, outer stamens plus carpel, inner stamens; or, sepals, petals, carpel, outer stamens, inner stamens. The outer stamen whorl and the carpel show possible overlap in time of initiation. Equalization of organ size occurs only within the stamen whorls. The sepals retain distinction in size, and the petals exhibit an inverse size to age relationship. The keel petals postgenitally fuse along part of their abaxial margins; their bases, however, remain free. Soybean flowers initiated at cool day/night temperatures of 18/14 C exhibited abnormalities and intermediate organs in all whorls. The gynoecium consisted of one to ten carpels (usually three or four), and carpel connation varied. Fusion of keel petals was often lacking, and stamen filaments fused erratically. Multiple carpellate flowers developed into multiple pods that were separate or variously connate. Intermediate type organs had characteristics only of organs in adjacent whorls. These aberrant flowers demonstrate that the floral meristem of soybean is not fixed or limited in its developmental capabilities and that it has the potential to produce alternate morphological patterns.     

荷花“重瓣化”的花器官形态发育比较观察

荷花（Nelumbo nucifera）的花型可分5种,最原始为单瓣型,然后由单瓣演化出半重瓣、重瓣、重台和千瓣型。为了揭示荷花重瓣化的分子机理,有必要从花器官的形态发育特征探究荷花花型成因及“重瓣化”的形态发育特征。实验分别选取5种荷花花型的代表品种：‘单洒锦’（单瓣型）、‘大洒锦’（重瓣型）、‘中山红台’（重台型）、‘至尊千瓣’（全重瓣型）、‘千瓣莲’（千瓣型）为材料,进行花芽分化过程形态的石蜡切片比较观察。结果发现：花芽分化过程中5个品种的萼片原基分化期和花瓣原基分化期相似,而雄蕊和雌蕊原基发育存在明显差异：单瓣、重瓣和重台品种均有正常的雄蕊和雌蕊原基分化;全重瓣品种发育初期有雄蕊及雌蕊原基分化,但后期全部瓣化;‘千瓣莲’品种不形成雄蕊和雌蕊原基,而是直接形成2至多个“花瓣增殖中心”,并由此不断分化出细小花瓣。研究认为重瓣型荷花品种的“重瓣化”花瓣主要来源于雄蕊的向心式瓣化,其次是雌蕊瓣化,属于雌雄蕊起源。而对于‘千瓣莲’型品种,花瓣的具体来源方式、花托是否直接参与瓣化及其在重瓣化过程中的作用有待于结合分子生物学手段开展进一步研究。     

大戟科麻疯树属三种植物花器官发生

利用扫描电子显微镜观察了大戟科Euphorbiaceae麻疯树属Jatropha麻疯树J. curcas L.、佛肚树J. podagrica Hook.和棉叶麻疯树J. gossypifolia L.花器官发生。结果表明: 麻疯树、佛肚树和棉叶麻疯树花萼原基均为2/5型螺旋发生。在同一个种不同的花蕾中, 花萼的发生有两种顺序: 逆时针方向和顺时针方向。远轴面非正中位的1枚先发生。5枚花瓣原基几乎同时发生。雄花中雄蕊两轮, 外轮对瓣, 内轮对萼。研究的3种麻疯树属植物雄蕊发生方式有两种类型: 麻疯树亚属麻疯树的5枚外轮雄蕊先同时发生, 5枚内轮雄蕊后同时发生, 佛肚树亚属佛肚树和棉叶麻疯树雄蕊8-9枚, 排成两轮, 内外轮雄蕊同时发生。雌花的3枚心皮原基为同时发生。麻疯树属单性花, 雌花的子房膨大而雄蕊退化, 雄花的雄蕊正常发育, 子房缺失。根据雄蕊发生方式, 支持将麻疯树属分为麻疯树亚属subgen. Jatropha和佛肚树亚属subgen. Curcas。     

Negative regulation of the Arabidopsis homeotic gene AGAMOUS by the APETALA2 product.

We characterized the distribution of AGAMOUS (AG) RNA during early flower development in Arabidopsis. Mutations in this homeotic gene cause the transformation of stamens to petals in floral whorl 3 and of carpels to another ag flower in floral whorl 4. We found that AG RNA is present in the stamen and carpel primordia but is undetectable in sepal and petal primordia throughout early wild-type flower development, consistent with the mutant phenotype. We also analyzed the distribution of AG RNA in apetela2 (ap2) mutant flowers. AP2 is a floral homeotic gene that is necessary for the normal development of sepals and petals in floral whorls 1 and 2. In ap2 mutant flowers, AG RNA is present in the organ primordia of all floral whorls. These observations show that the expression patterns of the Arabidopsis floral homeotic genes are in part established by regulatory interactions between these genes.     

Early flower development in Arabidopsis.

The early development of the flower of Arabidopsis thaliana is described from initiation until the opening of the bud. The morphogenesis, growth rate, and surface structure of floral organs were recorded in detail using scanning electron microscopy. Flower development has been divided into 12 stages using a series of landmark events. Stage 1 begins with the initiation of a floral buttress on the flank of the apical meristem. Stage 2 commences when the flower primordium becomes separate from the meristem. Sepal primordia then arise (stage 3) and grow to overlie the primordium (stage 4). Petal and stamen primordia appear next (stage 5) and are soon enclosed by the sepals (stage 6). During stage 6, petal primordia grow slowly, whereas stamen primordia enlarge more rapidly. Stage 7 begins when the medial stamens become stalked. These soon develop locules (stage 8). A long stage 9 then commences with the petal primordia becoming stalked. During this stage all organs lengthen rapidly. This includes the gynoecium, which commences growth as an open-ended tube during stage 6. When the petals reach the length of the lateral stamens, stage 10 begins. Stigmatic papillae appear soon after (stage 11), and the petals rapidly reach the height of the medial stamens (stage 12). This final stage ends when the 1-millimeter-long bud opens. Under our growing conditions 1.9 buds were initiated per day on average, and they took 13.25 days to progress through the 12 stages from initiation until opening.     

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.     

长角凤仙花（凤仙花科）的花器官发生和发育

以不同发育时期的长角凤仙花Impatiens longicornuta Y.L.Chen（凤仙花科Balsaminaceae）为材料,利用扫描电镜技术观察了其花器官的分化及其发育过程。长角凤仙花为两侧对称花,具2枚侧生萼片,唇瓣囊状,旗瓣具鸡冠状突起,雄蕊5枚,子房上位,5心皮5室。其花器官分化顺序为向心式,萼片—花瓣—雄蕊—雌蕊原基。2枚侧生萼片先发生,然后近轴萼片（即唇瓣）原基和2枚前外侧萼片原基近同时发生;但是这3枚萼片原基的发育不同步,远轴的2枚前外侧萼片原基的发育渐渐滞后,然后停止发育,最后渐渐为周围组织所吸收,直至消失不见。花瓣原基中,旗瓣原基最先发生,4个侧生花瓣原基相继成对发生,且之后在基部成对愈合形成翼瓣;5枚雄蕊原基几乎同时发生,5个心皮原基轮状同时发生。本文结果支持凤仙花属植物为5基数的花,并进一步证实了唇瓣的萼片来源;此外,研究结果表明花器官早期发育资料对植物系统与进化研究具有重要参考价值。     

荷花“重瓣化”的花器官形态发育比较观察

荷花（Nelumbo nucifera）的花型可分5种,最原始为单瓣型,然后由单瓣演化出半重瓣、重瓣、重台和千瓣型。为了揭示荷花重瓣化的分子机理,有必要从花器官的形态发育特征探究荷花花型成因及“重瓣化”的形态发育特征。实验分别选取5种荷花花型的代表品种：‘单洒锦’ （单瓣型）、‘大洒锦’ （重瓣型）、‘中山红台’ （重台型）、‘至尊千瓣’ （全重瓣型）、‘千瓣莲’ （千瓣型） 为材料,进行花芽分化过程形态的石蜡切片比较观察。结果发现：花芽分化过程中5个品种的萼片原基分化期和花瓣原基分化期相似,而雄蕊和雌蕊原基发育存在明显差异：单瓣、重瓣和重台品种均有正常的雄蕊和雌蕊原基分化;全重瓣品种发育初期有雄蕊及雌蕊原基分化,但后期全部瓣化;‘千瓣莲’品种不形成雄蕊和雌蕊原基,而是直接形成2至多个“花瓣增殖中心”,并由此不断分化出细小花瓣。研究认为重瓣型荷花品种的“重瓣化”花瓣主要来源于雄蕊的向心式瓣化,其次是雌蕊瓣化,属于雌雄蕊起源。而对于‘千瓣莲’型品种,花瓣的具体来源方式、花托是否直接参与瓣化及其在重瓣化过程中的作用有待于结合分子生物学手段开展进一步研究。     

Light and scanning electron microscopic studies of flower development inLindenbergia macrostachya Benth. (Scrophulariaceae)

All the floral primordia are homologous to leaves in their development inLindenbergia macrostachya. The sepals follow an anterior to posterior sequence of initiation. The petals and stamens are initiated almost simultaneously but sequentially in order of petals followed by stamens. There is no sign of development of fifth posterior stamen. p ]The calyx tube is formed by interprimordial growth followed by zonal growth. The combined interprimordial growth between the petal primordia and growth on the abaxial side of stamen primordia results in the formation of upper corolla tube whereas lower corolla tube is formed only by zonal growth. The zonal growth extends below the bases of stamen primordia also due to which they become epipetalous. The placentae arise from the carpellary margins, move inwards and get fused in the lower half and remain free in the upper part of the ovary. Thus the ovary appears biloeular with axile plaeentation in the lower haler and unilocular with parietal placentation in the upper half.     

泽苔草的花器官发生

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

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     

Inflorescence and floral development of Dahlstedtia species (Leguminosae: Papilionoideae: Millettieae)

Inflorescence and floral development of two tropical legume trees, Dahlstedtia pinnata and Dahlstedtia pentaphylla, occurring in the Atlantic Forest of south-eastern and southern Brazil, were investigated and compared with other papilionoids. Few studies have been made of floral development in tribe Millettieae, and this paper is intended to fill that gap in our knowledge. Dahlstedtia species have an unusual inflorescence type among legumes, the pseudoraceme, which comprises axillary units of three or more flowers, each with a subtending bract. Each flower exhibits a pair of opposite bracteoles. The order of flower initiation is acropetal; inception of the floral organs is as follows: sepals (5), petals (5), carpel (1) plus outer stamens (5) and finally inner stamens (5). Organ initiation in sepal, petal and inner stamen whorls is unidirectional; the carpel cleft is adaxial. The vexillum originates from a tubular-shaped primordium in mid-development and is larger than other petals at maturity, covering the keels. The filament tube develops later after initiation of inner-stamen primordia. Floral development in Dahlstedtia is almost always similar to other papilionoids, especially species of Phaseoleae and Sophoreae. But one important difference is the precocious ovule initiation (open carpel with ovules) in Dahlstedtia, the third citation of this phenomenon for papilionoids. No suppression, organ loss or anomalies occur in the order of primordia initiation or structure. Infra-generic differences in the first stages of ontogeny are rare; however, different species of Dahlstedtia are distinguished by the differing distribution pattern of secretory cavities in the flower.     

Original Paper Floral Organogenesis in Verbenaceae sensulato

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

Floral morphogenesis of Caltha and Trollius (Ranunculaceae) and its systematic significance

The floral morphogenesis of Caltha palustris L. and Trollius buddae Schipcz. was observed with a scanning electron microscope (SEM). The primordia of all floral organs initiate spirally and centripetally and develop centripetally. The spiral initiation sequence may be a basic pattern in Ranunculaceae. The primordia of bracts, sepals, and other floral organs are different in shape: the bract primordia are triangle, the sepal primordia crescent, and the petal (in Trollius), stamen, and carpel primordia hemispheric. This may indicate that the bracts, the sepals and other floral organs are different in origin. The petals are retarded in early developmental stages in Trollius buddae Schipcz, and have purses at the base. The retarded petals are very common in Ranunculaceae and the purse of the petal is similar to that of some Aquilegia species. The microspores in a longitudinal series of stamens develop centripetally in Caltha and Trollius; this may be a basic pattern in Ranunculaceae. The carpel primordia are plicate. In the developmental process of the carpels, the stigmatic tissue appears from the apex of the style and is decurrent along the ventral suture in Caltha, but there is no obvious stigmatic tissue in Trollius. Based on floral morphogenesis characteristics as well as the results from molecular systematics, comparative morphology and palynology studies, we consider that Caltha is not closely related to Trollius and that these two genera should not be treated in the same tribe.     

驴蹄草属和金莲花属(毛茛科)花器官的形态发生及系统学意义

利用扫描电镜观察了驴蹄草Caltha palustris L.和川陕金莲花Trollius buddae Schipcz.花器官的发生和发育过程。结果显示:驴蹄草和川陕金莲花的所有花器官均螺旋状向心式发生、向心式发育,花器官的螺旋状发生方式在毛茛科Ranunculaceae可能是一种基本式样;苞片、萼片与其他花器官原基的形状明显不同,显示苞片、萼片与其他花器官在系统发生上有所不同;川陕金莲花的花瓣在早期延迟发育且基部具囊,花瓣的延迟发育在毛茛科具花瓣的属中非常普遍,而花瓣基部的囊类似于耧斗菜属Aquilegia一些植物;两个属雄蕊群一纵列雄蕊中的小孢子均向心式发育,这种发育方式在毛茛科可能为基本类型。两个属植物的心皮原基均为对折式,在发育过程中,驴蹄草心皮顶端沿腹缝线形成下延的柱头组织,川陕金莲花不形成明显的柱头组织。根据花形态发生和发育特点,并结合其他研究成果,认为这两个属不应当属于同一个族。