商陆属植物的花器官发生

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

DIOECY IN BAUHINIA RESULTING FROM ORGAN SUPPRESSION

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

INITIATION OF STAMENS,CARPELS, AND RECEPTACLE IN THE CACTACEAE

Five taxa representing the three tribes of the Cactaceae have similar patterns of stamen and carpel initiation but display differences in early receptacle development. The first ring of stamens and the carpels arise simultaneously from subsurface layers. The bases of carpels are congenitally connate. Additional stamens are initiated centrifugally. The shape of the floral meristem within the ring created by the first stamens varies. In Pereskia corrugata it remains broadly convex; in Opuntia engelmannii it forms a depression with a small convex central region; in Epiphyllum strictum it forms a broad shallow depression; in Echinocereus reichenbachii var. albispinus it develops a deep depression; and in Mammillaria compressa it develops a depression prior to stamen and carpel initiation. Changes in receptacle shape result from cessation of apical growth and activation of an intercalary ring meristem. These two processes occur earlier in ontogeny in the more advanced of these five taxa.     

商陆属2种植物花器官发生过程观察

该研究采用扫描电镜观察红蕊商陆(Phytolacca esculenta)和浙江商陆(Phytolacca zhejiangensis)的花器官发生过程,以明确商陆属植物花的基数,以及雄蕊和雌蕊是否具有叶性器官发生的特点,阐明商陆属植物花发生的模式。观察结果显示:(1)红蕊商陆和浙江商陆在花原基发生后,小苞片以2/5圆周相继发生,花被片的发生紧接小苞片的发生进行,花被与小苞片的发生均有顺时针和逆时针方向,且二者的发生方向始终一致。(2)花被发生结束后,雄蕊在花顶端分生组织的环状分生组织上发生,没有明显的发生顺序,近似同时发生;2轮雄蕊时内轮雄蕊先发生;外轮雄蕊有少数有时偶然与花被互生,但因外轮雄蕊数多于花被数,雄蕊与花被常不互生,也没有规律性。(3)红蕊商陆和浙江商陆的心皮都在雄蕊发生后,紧接着开始发生,且雌蕊与雄蕊(或内轮雄蕊)互生发生;心皮没有发生的先后次序,且每个心皮在基部连成一个整体形成雌蕊基部并发育成为子房。(4)红蕊商陆和浙江商陆的花基数为5,雄蕊和雌蕊的发生及数目不符合5基数的特点。研究认为,红蕊商陆和浙江商陆为5基数花,该研究结果不支持商陆属植物为3基数花的发生模式。     

THE DEVELOPMENTAL BASIS FOR SEXUAL EXPRESSION IN CERATONIA SILIQUA (LEGUMINOSAE: CAESALPINIOIDEAE: CASSIEAE)

The flowers of Ceratonia siliqua, an anomalous caesalpinioid legume in the tribe Cassieae, are unusual in being unisexual and in lacking petals. Inflorescence development, organogeny, and flower development are described for this species. All flowers are originally bisexual, but one sex is suppressed during late development of functionally male and female flowers. Ceratonia siliqua is highly plastic in sexuality of individuals, inflorescence branching pattern, racemose or cymose inflorescences, bracteole presence, terminal flower presence, organ number per whorl, missing floral organs, pollen grain form, and carpel cleft orientation. Order of initiation is: five sepals in helical order, then five stamens in helical order together with the carpel. Each stamen is initiated as two alternisepalous primordia that fuse to become a continuous antesepalous ridge; in some flowers, the last one or two stamens of the five may form as individual antesepalous mounds. Petal rudiments are occasional in mature flowers. Position of organs is atypical: the median sepal is on the adaxial side in Ceratonia, rather than abaxial as in most other caesalpinioids. This feature in Ceratonia may be viewed as a link to subfamily Mimosoideae, in which this character state is constant.     

宽叶泽苔草的花器官发生

通过扫描电镜观察了宽叶泽苔草Caldesia grandisSamuel.的花器官发生。宽叶泽苔草 的萼片3枚,逆时针螺旋向心发生 ;花瓣3枚,呈一轮近同时发生,未观察到花瓣_雄蕊复合原基;雄蕊、心皮原基皆轮状向心 发生,最先近同时发生的6枚原基全部发育成雄蕊,随后发生的6枚原基早期并无差别,在发 育过程中逐渐出现形态差异,直至其中1－4枚发育成心皮,其余的发育成雄蕊;而后的几轮 心皮原基,6枚一轮,陆续向心相间发生。本文揭示了3枚萼片螺旋状的发生方式,并推测这种螺旋方式是泽泻科植物进化过程中保留下来     

Morphological studies in Zygophyllaceae. II. The floral development and vascular anatomy of Peganum harmala

Floral development and vascular anatomy are investigated in Peganum harmala, emphasizing its unusual androccium with 15 stamens. Sepals arise successively; petals emerge simultaneously with five antesepalous stamens. The five stamen pairs arise in the space between the petals and the antesepalous stamens. The gynoecium arises from three carpel primordia with evidence of two reduced carpels. Placentae are axile and each bears two double rows of ovules. A weakly developed nectary surrounds the base of the ovary. The antepetalous stamen traces diverge from a common supply to petals and sepal laterals, independent of the antesepalous stamen traces. The androecium of Peganum is described as a derived obdiploste-monous form, differing from the complex haplostemonous androecium of Nitraria. “Congenital dédoublement” cannot adequately explain the origin of the paired antepetalous stamens; two stamens can arise either by the splitting of a common primordium or independently, and both ways of inception are best understood as extremes of a gradation. The systematic position of Peganum is discussed in relation to other Zygophyllaceae using a cladistic analysis with Ptelea (Rutaceae) and Quassia (Simaroubaceae) as outgroups. The basal division in the Zygophyllaceae is between Peganum and the rest of the family.     

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

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

Floral development of petaloid Alismatales as an insight into the origin of the trimerous Bauplan in monocot flowers

Monocots are remarkably homogeneous in sharing a common trimerous pentacyclic floral Bauplan. A major factor affecting monocot evolution is the unique origin of the clade from basal angiosperms. The origin of the floral Bauplan of monocots remains controversial, as no immediate sister groups with similar structure can be identified among basal angiosperms, and there are several possibilities for an ancestral floral structure, including more complex flowers with higher stamen and carpel numbers, or strongly reduced flowers. Additionally, a stable Bauplan is only established beyond the divergence of Alismatales. Here, we observed the floral development of five members of the three ‘petaloid’ Alismatales families Butomaceae, Hydrocharitaceae, and Alismataceae. Outer stamen pairs can be recognized in mature flowers of Alismataceae and Butomaceae. Paired stamens always arise independently, and are either shifted opposite the sepals or close to the petals. The position of stamen pairs is related to the early development of the petals. In Butomaceae, the perianth is not differentiated and the development of the inner tepals is not delayed; the larger inner tepals (petals) only permit the initiation of stamens in antesepalous pairs. Alismataceae has delayed petals and the stamens are shifted close to the petals, leading to an association of stamen pairs with petals in so-called stamen–petal complexes. In the studied Hydrocharitaceae species, which have the monocot floral Bauplan, paired stamens are replaced by larger single stamens and the petals are not delayed. These results indicate that the origin of the floral Bauplan, at least in petaloid Alismatales, is closely linked to the position of stamen pairs and the rate of petal development. Although the petaloid Alismatales are not immediately at the base of monocot divergence, the floral evolution inferred from the results should be a key to elucidate the origin of the floral Bauplan of monocots.     

三白草科花部发育及其系统学意义

本研究从比较三白草科属间小花个体发育及分析花器官数量变异入手,探寻花器官在发生顺序、数目变化及排列方式等方面的演化趋势,揭示系统发育在个体发育中一定程度重现的事实及属间的进化关系。结果简述如下:首先,雄蕊和心皮发生顺序由中部优先演化到两侧优先。其次,由于远中雄蕊和心皮经历了从发育延迟、生长减缓到最终消失的历程,中部雄蕊和心皮由成对演化为单生。此外,两侧生雄蕊对由各自独立的原基发生演化到共同原基发生或减化为1枚,假银莲花属近中1枚雄蕊原基二裂成1对,蕺菜属3枚心皮发生于一环状共同原基等,都是该科花器官演化的重要事实并可归结为融合、减化和复化的结果。文章根据花器官的演化趋势及过渡类型的剖析,论述了三白草科属间的系统进化关系。     

月季花瓣数量遗传分析

以‘窄叶藤本月季花’( Rosa chinensis ‘Zhaiye Tengben Yuejihua’)ב月月粉’( R. chinensis ‘Old Blush’)杂交群体为材料, 分析其花瓣数量的分离特点, 对单瓣花与重瓣花的花芽分化过程进行观察, 并对花瓣、雄蕊及瓣化雄蕊进行表皮细胞超微结构的观察.结果显示...     

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。     

本地种西番莲与外引种西番莲的花外蜜腺-蚂蚁-植食昆虫相互作用系统的比较

外引植物如何和当地生物建立相互关系是入侵生物学和进化生物学关注的关键科学问题.本实验以西双版纳当地自然分布的长叶西番莲(Passiflora siamica)和近年从南美新引种的红花两番莲(P.coccinea)为材料,探讨两种西番莲植物-蚂蚁-植食昆虫间相互关系系统对变化响应的差异.结果表明,共有24种蚂蚁取食花外蜜液,利用两种西番莲的蚂蚁数量没有显著差异(t=1.30,P=0.20),但在种类组成上存在显著差异(χ~2=14.76,df=4,P<0.01).植食昆虫多为蜗牛、螽斯、鳞翅目幼虫等广适性昆虫.去除蚂蚁处理均未对两种西番莲植物的平均叶面积损失造成显著影响.该研究表明两种两番莲的花外蜜腺-蚂蚁-广适性昆虫的相互作用系统对变化的响应没有显著差异,而与专食性昆虫相互作用系统对变化的响应的差异则有待进一步研究.     

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.     

Floral ontogeny in legume genera Petalostylis, Labichea, and Dialium (Caesalpinioideae: Cassieae), a series in floral reduction

Floral ontogeny of taxa of two subtribes (Labicheinae, Dialiinae) of caesalpinioid tribe Cassieae, characterized by reduced number of floral organs, was compared. All three taxa studied are distichous; Petalostylis labicheoides flowers are solitary in leaf axils, Labichea lanceolata has few-flowered racemes, and Dialium guineense has numerous-flowered cymes. The first sepal primordium in each is initiated abaxially and nonmedianly. Order of organogenesis in Petalostylis is: five sepals bidirectionally, five petals and carpel simultaneously, then five stamens bidirectionally, starting abaxially. The order in Labichea is: five sepals helically (one lagging in time), five petals unidirectionally starting abaxially, the carpel and petals concurrently, then two stamens successively, starting laterally. Order in Dialium is: five sepals bidirectionally, the single petal adaxially, and lastly the carpel and two stamens concurrently. Specializations include (1) reduction of the five sepals to four by fusion in Petalostylis and Labichea; (2) reduction of petal number to one in Dialium; (3) reduction of stamen number to two in Labichea and Dialium, and reduction of functional stamens to three in Petalostylis; and (4) an elaborate, late-developing style in Petalostylis. Floral asymmetry, another specialization, characterizes Labichea, expressed by dissimilar stamens, while the other genera have zygomorphic flowers. Floral ontogenies are compared with other taxa of Cassieae.     

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.     

The discovery of polyandry in Curculigo (Hypoxidaceae): implications for androecium evolution of asparagoid monocotyledons

BACKGROUND AND AIMS: Individual flowers of the monocot Curculigo racemosa (Hypoxidaceae, Asparagales) are regularly polyandrous. To evaluate the significance of this almost unique character among Asparagales for flower evolution of asparagoid monocots, flowers of C. racemosa were studied comparatively. METHODS: Anthetic flowers as well as early floral developmental stages were studied by light and scanning electron microscopy. KEY RESULTS: Despite the polyandry, floral development is similar to that of other Asparagales with a developmental gradient from adaxial to abaxial. Stamens initiate simultaneously and the diameter of staminal primordia is about half of that in species with six anthers. The number of stamens is not fixed (12-26) and varies within the same inflorescence. Surprisingly, the gynoecium can be four- or six-locular, besides the normal trimerous state. CONCLUSIONS: The discovery of a polyandrous Curculigo reveals plasticity of stamen number at the base of Asparagales. Orchidaceae - sister to all other Asparagales - has a reduced stamen number (three, two or one), whereas in Hypoxidaceae - part of the next diverging clade - either the normal monocot stamen number (six), polyandry (this study) or the loss of three anthers (Pauridia) occurs. However, at present it is impossible to decide whether the flexibility in stamen number is autapomorphic for each group or whether it is a synapomorphy. The small size of stamen primordia of Curculigo is conspicuous. It allows more space for additional androecial primordia. Stamens are initiated as independent organs, and filaments are not in bundles, hence C. racemosa is not secondarily polyandrous as may be the case in the distantly related Gethyllis of asparagoid Amaryllidaceae. The increase in carpel number is a rare phenomenon in angiosperms. A possible explanation for the polyandry of C. racemosa is that it is a natural SUPERMAN-deficient mutant, which shows an increase of stamens, or ULTRAPETALA or CARPEL FACTORY mutants, which are polyandrous and changed in carpel number.     

Androecium and floral nectaries ofHarungana madagascariensis (Clusiaceae)

The mature flower ofHarungana madagascariensis (Choisy)Poir. has an androecium of five antipetalous fascicles, consisting of four stamens each. The stamen fascicles alternate with five indented nectary scales. A SEM-study of the floral development, as well as a study of the floral anatomy was carried out to understand whether the nectariferous scales represent staminodia or are receptacular in nature and consequently whether or not the androecium ofHarungana, and theClusiaceae in general, is originally diplostemonous. The five petals originate by the splitting of petal-stamen complexes. Next the upper part of each complex differentiates basipetally in four stamens. The stamens remain fascicled and are lifted on a long stalk at maturity. Five carpel primordia are initiated united in a low ringwall. The five nectary scales appear after carpel inception and develop an external morphology reminiscent of anthers. The floral anatomy reveals an independent origin of sepal median traces and common sepal lateral traces, free petal traces, stamen fascicle traces and alternating vascular tissue which supplies the nectaries. The petal-stamen complexes are the result of a retardation in petal inception, linked with the absorption of petal tissue into the stamen primordia. The development of the stamen fascicles is discussed; it is suggested that they are of a secondary nature and do not appear as a reduction from a multistaminate androecium. The external morphology and vascular anatomy of the scales speaks in favour of a staminodial nature. The comparison with some other species of theClusiaceae gives evidence of a diplostemonous ancestry of the androecium.     

Floral ontogeny of <Emphasis Type="Italic">Schisandra chinensis</Emphasis> (Schisandraceae): implications for androecial evolution within <Emphasis Type="Italic">Schisandra</Emphasis> and <Emphasis Type="Italic">Kadsura</Emphasis>

The organogenesis of staminate and carpellate flowers of Schisandra chinensis (Schisandraceae) was investigated with scanning electron microscopy, with observations on the development of tepals reported for the first time. The results showed that there is no interval between the initiation of the last tepal and that of the first stamen or carpel, and that the shapes of tepal, stamen, and carpel primordia are similar. The tepals and stamens of staminate flowers are initiated acropetally in a continuous spiral Fibonacci phyllotaxis, with no carpel structures observed; the filaments are not connate. The organogenesis of the carpellate flowers is similar to that of the staminate flowers, but with no evidence of stamen development. The carpels are ascidiate without postgenital fusion. Three androecial characters of Schisandra and Kadsura are discussed in a phylogenetic context. The subglobose or obovoid androecium of Schisandra propinqua and Schisandra plena may be homologous with that in sections Kadsura and Sarcocarpon. The plesiomorphic form of the androecium within the two genera is likely to be elongate with more than ten free stamens.