Meristic variation in Potamogeton flowers

Flowers of Potamogeton normally have a completely tetramerous plan. Deviations from this norm occur quite commonly in the uppermost flowers of the inflorescence; these variations have been reported before and usually involve a reduction in number of parts. Cases have now been found where the gynoecium of all or many flowers differs from the normal tetracarpellate arrangement; some species regularly have fewer and others more than four carpels. The developmental bases of meristic variation have been explored and quantitative studies of gynoecia and developing gynoecia have been undertaken. The data are used to evaluate the control and correlation of floral development in Potamogeton in general, and in particular the relationship between the gynoecium and the rest of the flower. The developing flower passes through two successive phases of organ initiation: one in which the perianth and stamen primordia arise, and one in which the gynoecial primordia arise. There seems to be little developmental relationship between the two phases except phyllotactic continuity. During the perianth/stamen phase each stamen primordium arises directly above a perianth member, and the presence of a perianth member seems to be a prerequisite for initiation of the stamen. The perianth/stamen phase seems to be rather stable so that normally four perianth/stamen associations are initiated, except in flowers at the tip of the inflorescence. In the gynoecial phase the number of carpel primordia initiated seems to depend on the relative size of carpel primordia and floral apex, and on whether or not the floral apex continues to grow while initiating carpel primordia.     

Developmental study on the inflorescence and flower of Caldesia grandis Samuel (Alismataceae)

The inflorescence and floral development of Caldesia grandis Samuel is reported for the first time in this paper. The basic units of the large cymo‐thyrsus inflorescence are short panicles that are arranged in a pseudowhorl. Each panicle gives rise spirally to three bract primordia also arranged in a pseudowhorl. The branch primordia arise at the axils of the bracts. Each panicle produces spirally three bract primordia with triradiate symmetry (or in a pseudowhorl) and three floral primordia in the axils of the bract primordia. The apex of the panicle becomes a terminal floral primordium after the initiations of lateral bract primordia and floral primordia. Three sepal primordia are initiated approximately in a single whorl from the floral primordium. Three petal primordia are initiated alternate to the sepal primordia, but their subsequent development is much delayed. The first six stamen primordia are initiated as three pairs in a single whorl and each pair appears to be antipetalous as in other genera of the Alismataceae. The stamen primordia of the second whorl are initiated trimerously and opposite to the petals. Usually, 9–12 stamens are initiated in a flower. There is successive transition between the initiation of stamen and carpel primordia. The six first‐initiated carpel primordia rise simultaneously in a whorl and alternate with the trimerous stamens, but the succeeding ones are initiated in irregular spirals, and there are 15–21 carpels developed in a flower. Petals begin to enlarge and expand when anthers of stamens have differentiated microsporangia. Such features do not occur in C. parnassifolia. In the latter, six stamen primordia are initiated in two whorls of three, carpel primordia are initiated in 1–3 whorls, and there is no delay in the development of petals. C. grandis is thus considered more primitive and C. parnassifolia more derived. C. grandis shares more similarities in features of floral development with Alsma, Echinodorus, Luronium and Sagittaria. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140 , 39–47.     

PERIANTH DEVELOPMENT OF POTAMOGETON RICHARDSONII

Interpretation of the Potamogeton flower is complicated by the attachment of the “perianth segment” to the stamen connective. Developmental studies show that the perianth segments are not outgrowths of the stamen connectives. They are initiated on the floral apex acropetally before the (superposed) primordia of the stamens. After the inception of the stamen primordia, growth occurs in the regions between the primordia of each perianth segment and stamen. Thereby the bases of the developing perianth segment and stamen become united, and in the adult flower eventually the perianth segment is inserted on the connective of the stamen. The primordium of the perianth segment develops from the 2 outer layers (tunica) of the floral apex, in contrast to the stamen primordium which originates from the 3 outer layers. The vascular bundles for each perianth segment–stamen region develop acropetally from 1 common bundle which bifurcates into 1 bundle for the perianth segment and 1 for the stamen. The bundle leading into the perianth segment branches in a more or less dichotomous manner. The veins form none or only 1 or 2 anastomoses at the base of the lamina, whereas the vein endings remain free. The interpretation of the perianth segments is discussed in terms of the classical and the gonophyll theory. Since both theories rest on an ambiguous methodological basis, interpretation is postponed until a new approach to comparative morphology has been worked out and until the floral development of other Helobiales has been studied.     

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.     

灌木铁线莲（毛茛科）花器官的发生与发育

用扫描电子显微镜（SEM）对铁线莲属（Clematis L.）植物灌木铁线莲(C. fruticosa Turcz.)花的形态发生和发育过程进行了观察。灌木铁线莲花原基形成后,4枚萼片以交互对生的方式首先发生,呈轮状排列。最早的4枚雄蕊原基在4枚萼片交接的位置上近螺旋状发生,此后,随着雄蕊原基的向心发生和数目不断增多,其发生的螺旋状序列逐渐明显。雄蕊原基发生后,在花原基顶端,心皮原基沿着雄蕊原基的发生序列呈螺旋状发生。本文结果支持在原始被子植物花中螺旋状排列和轮状排列同时存在的观点。此外,本文也进一步证实了花萼与苞片的同源性。     

Elucidating the unusual floral features of Swartzia dipetala (Fabaceae)

In this study, we evaluated the floral ontogeny of Swartzia dipetala, which has peculiar floral features compared with other legumes, such as an entire calyx in the floral bud, a corolla with one or two petals, a dimorphic and polyandrous androecium and a bicarpellate gynoecium. We provide new information on the function of pollen in both stamen morphs and whether both carpels of a flower are able to form fruit. Floral buds, flowers and fruits were processed for observation under light, scanning and transmission electron microscopy and for quantitative analyses. The entire calyx results from the initiation, elongation and fusion of three sepal primordia. A unique petal primordium (or rarely two) is produced on the adaxial side of a ring meristem, which is formed after the initiation of the calyx. The polyandrous and dimorphic androecium also originates from the activity of the ring meristem. It produces three larger stamen primordia on the abaxial side and numerous smaller stamen primordia on the adaxial side. These two types of stamens bear morphologically similar ripening pollen grains. However, prior to the dehiscence of thecae and presentation of pollen in the anther, only the pollen grains of the larger stamens contain amyloplasts. Two carpel primordia are initiated as distinct protuberances, alternating with the larger stamens, in a slightly inner position in the floral meristem, constituting the bicarpellate gynoecium. Both carpels are able to form fruit, although only one fruit is generally produced in a flower. The increase in gynoecium merism probably results in an increase in the surface deposition of pollen grains and consequently in the chance of pollination. This is the first study to thoroughly investigate organogenesis and the ability of the carpel to form fruit in a bicarpellate flower from a member of Fabaceae, in addition to the pollen ultrastructure in the heteromorphic stamens associated with the ‘division of labour’ sensu Darwin. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 303–320.     

The origin and evolution of carpels and fruits from an evo-devo perspective

The flower is an evolutionary innovation in angiosperms that drives the evolution of biodiversity.The carpel is integral to a flower and develops into fruits after fertilization,while the perianth,consisting of the calyx and corolla,is decorative to facilitate pollination and protect the internal organs,including the carpels and stamens.Therefore,the nature of flower origin is carpel and stamen origin,which represents one of the greatest and fundamental unresolved issues in plant evolutionary bi...     

Floral morphogenesis in Euptelea (Eupteleaceae, Ranunculales)

BACKGROUND AND AIMS: Based on molecular phylogenetic studies, the unigeneric family Eupteleaceae has a prominent phylogenetic position at or near the base of Ranunculales, which, in turn, appear at the base of eudicots. The aim of the present paper is to reveal developmental features of the flowers and to put the genus in a morphological context with other basal eudicots. METHODS: Flowers in all developmental stages of Euptelea pleiosperma were collected in the wild at intervals of 7-10 d in the critical stages and studied with a scanning electron microscope. KEY RESULTS: Remnants of a perianth are lacking throughout flower development. Floral symmetry changes from monosymmetric to asymmetric to disymmetric during development. Asymmetry is expressed in that the sequence of stamen initiation is from the centre to both lateral sides on the adaxial side of the flower but starting from one lateral side and proceeding to the other on the abaxial side. Despite the pronounced floral disymmetry, a dimerous pattern of floral organs was not found. The carpel primordia arise between the already large stamens and alternate with them. Stamens and carpels each form a somewhat irregular whorl. The carpels are ascidiate from the beginning. The stigma differentiates as two crests along the ventral slit of the ovary. The few lateral ovules alternate with each other. CONCLUSIONS: Although the flowers have some unusual autapomorphies (wind pollination, lack of a perianth, pronounced disymmetry of the floral base, long connective protrusion, long temporal gap between androecium and gynoecium initiation, small space for carpel initiation), they show some plesiomorphies at the level of basal eudicots (free carpels, basifixed anthers, whorled phyllotaxis), and thus fit well in Ranunculales.     

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 DEVELOPMENT OF POTAMOGETON RICHARDSONII

The inception and development of the sterile floral appendages of Potamogeton richardsonii have been re-investigated with a refined dissection technique (Sattler, 1968) and improved microtechnical methods (Feder and O'Brien, 1968). The results obtained by Sattler (1965) are confirmed, i.e., the sterile appendages are initiated at the flanks of the floral apex before the stamen primordia are formed. Consequently, they may be homologized with tepals or perianth members, although in the mature flower they are inserted at the stamen connective, due to growth between and at the base of each developing tepal and stamen. Each carpel arises as a radial primordium which becomes peltate immediately after its inception. One ovule primordium is initiated at the cross-zone. The stigma becomes bilobed. A slight outgrowth develops at the abaxial side of the style. The floral apex has a two-layered tunica. The primordia of the tepals, carpels, and ovules arise by periclinal divisions in the second tunica layer, whereas the stamen primordia are initiated by periclinal divisions in the corpus and second tunica layer. Variation in floral pattern, especially with regard to the number of appendages, has been observed in flowers near the tip of the inflorescence axis.     

FLORAL DEVELOPMENT IN GYMNOTHECA CHINENSIS (SAURURACEAE)

The development of the inflorescence and flowers are described for Gymnotheca chinensis Decaisne (Saururaceae), which is native only to southeast China. The inflorescence is a short terminal spike of about 50–70 flowers, each subtended by a small bract. There are no showy involucral bracts. The bracts are initiated before the flowers, in acropetal order. Flowers tend to be initiated in whorls of three which alternate with the previous whorl members. No perianth is present. The flower contains six stamens, and four carpels fused in an inferior ovary containing 40–60 ovules on four parietal placentae. Floral symmetry is dorsiventral from inception and throughout organ initiation. Floral organs are initiated in the following order: 1) median adaxial stamen, 2) a pair of lateral common primordia which bifurcate radially to produce two stamen primordia each, 3) median abaxial stamen, 4) a pair of lateral carpel primordia, 5) median adaxial carpel, 6) median abaxial carpel. This order of initiation differs from that of any other Saururaceae previously investigated. The inferior ovary results from intercalary growth below the level of stamen attachment; the style elongates by intercalary growth, and the four stigmas remain free. The floral structure of Gymnotheca is relatively advanced compared to Saururus, but its assemblage of specializations differs from that of either Anemopsis or Houttuynia, the other derived genera in the Saururaceae.     

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。     

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 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.     

Morphology and development of the reproductive shoots of Lilaea scilloides (Poir.) Hauman (Alismatidae)

Shoots of adult plants of Lilaea scilloides have a sympodial form. Each unit of the sympodium bears a single sheathing prophyll (which is the only kind of foliage leaf produced in the adult) and terminates in an inflorescence. The prophyll subtends the next unit of the sympodium. A further accessory bud can form in association with each unit. This bud repeats the pattern of the main sympodium, giving the plant a tufted habit. Five different kinds of flower can be identified in the inflorescence: a unisexual male flower with a single perianth member and adnate stamen; a bisexual flower, with a single perianth member and adnate stamen, and a single carpel with an anatropous bitegmic ovule; a unisexual female flower with a single perianth member and carpel; a unisexual female flower comprising only a single carpel; and a female flower comprising only a single carpel with a very long filamentous style. The first four kinds occur in the upper part of the inflorescence which is normally elevated on a scape, while the last kind is restricted to the base of the inflorescence. In the position of the basal flowers several variations have been observed in cultivated material. These include branching associated with the basal flowers, which results in the development of additional basal flowers or inflorescences, and even total replacement of a basal flower by an inflorescence or a branching structure bearing flowers. A review of past literature includes a clarification of some persistent errors which have confused the taxonomic position of the plant and the morphological interpretation of the reproductive appendages.     

Male and female flowers of the dioecious plant sorrel show different patterns of MADS box gene expression.

Male and female flowers of the dioecious plant sorrel (Rumex acetosa) each produce three whorls of developed floral organs: two similar whorls of three perianth segments and either six stamens (in the male) or a gynoecium consisting of a fertile carpel and two sterile carpels (in the female). In the developing male flower, there is no significant proliferation of cells in the center of the flower, in the position normally occupied by the carpels of a hermaphrodite plant. In the female flower, small stamen primordia are formed. To determine whether the organ differences are associated with differences in the expression of organ identity genes, cDNA clones representing the putative homologs of B and C function MADS box genes were isolated and used in an in situ hybridization analysis. The expression of RAD1 and RAD2 (two different DEFICIENS homologs) in males and females was confined to the stamen whorl; the lack of expression in the second, inner perianth whorl correlated with the sepaloid nature of the inner whorl of perianth segments. Expression of RAP1 (a PLENA homolog) occurred in the carpel and stamen whorls in very young flower primordia from both males and females. However, as soon as the inappropriate set of organs ceased to develop, RAP1 expression became undetectable in those organs. The absence of expression of RAP1 may be the cause of the arrest in organ development or may be a consequence.     

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.     

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     

掌叶木的花器官发生及其系统学意义

利用扫描电子显微镜和光学显微镜观察了掌叶木的花器官发生过程。观察结果表明: 花序原基最先发生, 然后形成两个大小不一的花原基; 萼片原基的发生不同步, 螺旋状向心发生; 4-5枚花瓣原基以接近轮状方式近同时发生; 不存在花瓣-雄蕊复合原基; 7-8枚雄蕊原基为近同时发生, 其生长较花瓣原基快; 心皮原基最后发生, 3枚心皮原基为同时发生。花为单性花。在雌花中, 子房膨大而雄蕊退化。在雄花中, 雄蕊正常发育, 子房退化。讨论了掌叶木花器官发生和发育的系统学意义。     

PATTERNS OF FLORAL DEVELOPMENT IN AGALINIS AND ALLIES (SCROPHULARIACEAE). II. FLORAL DEVELOPMENT OF AGALINIS DENSIFLORA

Initiation of floral primordia begins in Agalinis densiflora with production of two lateral adaxial calyx lobe primordia followed by a midadaxial primordium, and then primordia of two abaxial calyx lobes. Initiation of three abaxial corolla lobe primordia is succeeded by that of two stamen pairs and then by primordia of two adaxial corolla lobes. The primordium of the abaxial carpel appears before the adaxial one. Except for the calyx, initiation of primordia proceeds unidirectionally from the abaxial to the adaxial side of the floral apex. Zygomorphy in the calyx, corolla, and androecium is evident during initiation of primordia and is accentuated during organogenesis. The calyx undergoes comparatively rapid organogenesis, but the inner three floral series undergo a protracted period of organogenesis. The perianth series reach maturation prior to meiosis in the anthers. Maturation of the androecium and gynoecium are postmeiotic events.