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.     

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.     

Morphology of the gynoecium and systematic position of the Ochnaceae

The gynoecium is syncarpous in all Ochnaceae. In the Ochnoideae carpels are peltate with a conventional cross-zone bearing one ovule, or, in Lophira , a very broad cross-zone with an horizontal ovular row. In Ochna and Brackenridgea , the style is gynobasic, each carpel develops transmitting tissue on its morphologically dorsal surface, and this tissue lines a canal or originates a solid inner strand in each carpel at style level. The style is tubular, with an inner cuticle, and compound, each component with its own transmitting tissue. In Ouratea the style is solid with a single compound transmitting strand. In Lophira and Elvasia the transmitting tissue seems to be developed by the morphologically ventral carpellary surfaces. Ovules are unitegmic with a bivalent integument.
In the Sauvagesioideae carpels are peltate, but with ovules above the cross-zones, on margins of the symplicate zone. In Euthemis , there is one ovule on each side of, and close to, each cross-zone. The single stylar canal is bounded by the morphologically dorsal carpellary surfaces. In Sauvagesia ovules occur on both sides of the cross-zones but most of them are above on carpel margins, as are all ovules of Cespedesia. The stylar canal of Sauvagesia is bounded by the ventral carpel surfaces, three strips of the outer surface passing inside at the sutures and developing into transmitting tissue. The stylar canal of Cespedesia is bounded by the dorsal carpel surfaces. The gynoecium of Wallacea has two epeltate carpels with a laminar placentation, the carpel margins being displaced on to the topographically ventral carpel surfaces with a row of ovules along each margin. Ovules are bitegmic.
The Ochnoideae, which shows relationships with the Rutaceae, Meliaceae, Simaroubaceae and Hippocastanaceae, is more advanced than the Sauvagesioideae, which clearly belongs in the Violales. The Ochnaceae is to be placed in the Violales.     

COMPARATIVE ONTOGENY OF THE INFLORESCENCE AND FLOWER OF HAMAMELIS VIRGINIANA AND LOROPETALUM CHINENSE (HAMAMELIDACEAE)

A comparative developmental study of the inflorescence and flower of Hamamelis L. (4-merous) and Loropetalum (R. Br.) Oliv. (4–5 merous) was conducted to determine how development differs in these genera and between these genera and others of the family. Emphasis was placed on determining the types of floral appendages from which the similarly positioned nectaries of Hamamelis and sterile phyllomes of Loropetalum have evolved. In Hamamelis virginiana L. and H. mollis Oliv. initiation of whorls of floral appendages occurred centripetally. Nectary primordia arose adaxial to the petals soon after the initiation of stamen primordia and before initiation of carpel primordia. In Loropetalum chinense (R. Br.) Oliv. floral appendages did not arise centripetally. Petals and stamens first arose on the adaxial portion, and then on the abaxial portion of the floral apex. The sterile floral appendages (sterile phyllomes of uncertain homology) were initiated adaxial to the petals after all other whorls of floral appendages had become well developed. In all three species, two crescent shaped carpel primordia arose opposite each other and became closely appressed at their margins. Postgenital fusion followed and a falsely bilocular, bicarpellate ovary was formed. Ovule position and development are described. The nectaries of Hamamelis and sterile phyllomes of Loropetalum rarely develop as staminodia, suggesting a staminodial origin. However, these whorls arise at markedly different times and are therefore probably not derived from the same whorl of organs in a common progenitor. This hypothesis seems probable when one considers that the seemingly least specialized genus of the tribe, Maingaya, bears whorls of both staminodia and sterile phyllomes inside its whorl of stamens.     

Developmental Morphology and Histogenesis of Reproductive Structures of the Millet,Panicum miliaceum L. Histogenesis of the Inflorescence and Flower

The branches of successive orders of the inflorescence of Panicum miliaceum L. arise in the axils of the bracts of the branches of next lower order. Their initiation is evidenced by periclinal division of sub-hypodermal cells. The primordia of branches arise in initiation like a normal axillary bud. The floral histogenesis of Panicum miliaceum L. is similar to that of Triticum. Primordia of the spikelet, flower and stamen are initiated by the activity of the periclinal division of the sub-hypodermal cell or cells. Sometimes, periclinal divisions also occur in a few hypodermal cells during these primordial developments; such divisions are more frequent in the formation of the flower and stamen primordia than in the formation of the spikelet primordia. The periclinal division of the dermatogen ceils never occurs in the formation of these organs. Glumes and lemma are initiated in the periclinal division of the dermatogen and hypodermal cell or cells. The primordia of the palea, lodicule and carpel are initiated by means of the periclinal division in the dermatogen cell or cells. In the formation of the palea and carpel, periclinal divisions also occur in hypodermat cells, but their derivatives are protruding into the bases of the primordia and do not constitute the tissues of the palea and carpel. The growing point of the flower axis develops into the ovule. The integuments arise from the periclinal division of dermatogen cells. The periclinal division of dermatogen cells is characteristic of the initiation of the phylloid organs in the Gramineae.     

Floral ontogeny inMazus pumilus (Scrophulariaceae)

InMazus pumilus, all the floral appendages are initiated in acropetal sequence in the second cell layer (except stamens) of the floral primordium by periclinal divisions. The actinomorphic calyx tube is formed due to zonal growth. The zygomorphy in corolla is evident from the inception of petal primordia which arise sequentially as independent units in order of one anterior, a pair of anterio-lateral followed by a pair of posterio-lateral. Later these primordia exhibit differential growth because of which zygomorphy becomes more pronounced. The upper corolla tube is formed by interprimordial growth and lower corolla tube by zonal growth. Stamens are initiated in the third layer of the floral apex. Unlike sepals and petals, in the development of stamens (4) underlying cells of corpus also contribute. Posterior stamen is absent. The stamens become epipetalous because of interprimordial and zonal growth in the common region below the bases of petals as well as stamens. The two carpel primordia arise as crescent shaped structures which become continuous due to interprimordial growth. The ovary is formed by a ring of zonal meristem. The style develops later between stigma and ovary because of intercalary growth. The residual apex grows vertically along with the ovary and forms the septum of the ovary. All the floral appendages exhibit similar pattern of histogenesis and early growth suggesting thereby the appendicular nature of these appendages.     

MADS-box protein complexes control carpel and ovule development in Arabidopsis

The AGAMOUS (AG) gene is necessary for stamen and carpel development and is part of a monophyletic clade of MADS-box genes that also includes SHATTERPROOF1 (SHP1), SHP2, and SEEDSTICK (STK). Here, we show that ectopic expression of either the STK or SHP gene is sufficient to induce the transformation of sepals into carpeloid organs bearing ovules. Moreover, the fact that these organ transformations occur when the STK gene is expressed ectopically in ag mutants shows that STK can promote carpel development in the absence of AG activity. We also show that STK, AG, SHP1, and SHP2 can form multimeric complexes and that these interactions require the SEPALLATA (SEP) MADS-box proteins. We provide genetic evidence for this role of the SEP proteins by showing that a reduction in SEP activity leads to the loss of normal ovule development, similar to what occurs in stk shp1 shp2 triple mutants. Together, these results indicate that the SEP proteins, which are known to form multimeric complexes in the control of flower organ identity, also form complexes to control normal ovule development.     

FLORAL ONTOGENY OF ILLICIUM FLORIDANUM,WITH EMPHASIS ON STAMEN AND CARPEL DEVELOPMENT

The ontogeny of the flower and fruit of Illicium floridanum Ellis, the Star Anise, was investigated. Each of 5 or 6 bracts in each mixed terminal bud subtends either a vegetative or floral bud. The solitary flowers occur in terminal or axillary positions. Each flower has 3–6 subtending bracteoles arranged in a clockwise helix. The flowers in our material have 24–28 tepals, 30–39 stamens, and usually 13 (rarely 19) uniovulate carpels. Tepals and stamens are initiated in a low-pitched helix; carpels later appear whorled, but arise successively at different levels on the apical flanks. The floral apex is high-convex in outline with a tunica-corpus configuration; it increases in height and width throughout initiation of the floral appendages. Tepals, stamens, and carpels are initiated by one to several periclinal divisions in the subsurface layers low on the apical flanks, augmented by cell divisions in the outer layers of the corpus. The carpel develops as a conduplicate structure with appressed, connivent margins. Procambium development of floral appendages is acropetal and continuous. Bracteoles, tepals, stamens and carpels are each supplied by 1 trace; the carpellary trace splits into a dorsal and an ascending ventral sympodium. The latter bifurcates to form 2 ventral bundles. The ovular bundle diverges from the ventral sympodium. Ovule initiation occurs in a median axillary position to the carpel, an unusual type of ovule initiation. The fruit vasculature is greatly amplified as the receptacle and follicles enlarge. After carpel initiation an apical residuum persists which is not vascularized; a plate meristem develops over its surface to produce a papillate structure.     

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。     

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     

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.     

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。     

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.     

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.     

Gynoecial morphology and funicular germination plugs in the Nolanaceae

In Nolana humifusa (Gouan) Johnst. and N. paradoxa Lindl, five carpel primordia unite by their thin margins to form the gynoecium wall. An ovule primordium is initiated in each of a maximum of six depressions, formed in the adaxial surface of each carpel primordium. The depressions become deeper, each developing into a duct that ends in an ovule chamber, which is a uniovulate locellus. The locellus is delimited by a ventral carpellary epidermis except at its lower adaxial part, where the ovule is invaginated on a short funicle from its own placenta.
Periclinal cell divisions in the subsurface layers of the floral apex form a receptacular column, which grows in continuity with the lower adaxial parts of the carpel primordia; the upper parts of the carpel primordia face the five-radiate "common cavity" inside the gynoecium wall. At anthesis this cavity is filled with stylar and ovarian transmitting tissue. The latter forms five "wings" that downward are continuous with wings of the receptacular column and which together with them radiate between the five carpels. True septa are not formed.
In N. humifusa a plurilocellate mericarp originates from each carpel primordium. In N. paradoxa longitudinal unilocellate portions of each plurilocellate carpel primordium develop independently into "carpel-lobes", the bulging lower parts of which mature into unilocellate mericarps. In both species the funicle develops into a germination plug.
The locellar organization described is a common feature of the Nolanaceae. The formation of invariably uniovulate placentae in pluriovulate carpels is the basic innovation of the family.     

FLORAL ORGANOGENESIS IN NICOTIANA TABACUM: A COMPARISON OF TWO CYTOPLASMIC MALE-STERILE CULTIVARS WITH A MALE-FERTILE CULTIVAR

Flower buds of two cytoplasmically male-sterile cultivars and one male-fertile cultivar of Nicotiana tabacum (L.) were studied by light and scanning electron microscopy to compare their floral ontogenies. In [gla]tbc, a cultivar with N. tabacum nuclear genes and N. glauca cytoplasm, the five small stamen primordia cease growth soon after they are initiated. They remain unchanged at the flanks of the ovary throughout flower maturation, so the mature flower appears stamenless. In `[pbg]tbc,‘ a N. tabacum cultivar with cytoplasm that was thought to have been derived from N. plumbaginifolia, the divergence from normal stamen development occurs before carpel emergence at which point the primordia are already notably smaller and less cylindrical than primordia of the male-fertile cultivar. After carpel emergence the primordia continue to develop, forming pink structures called petalodes, which are more similar to petals than stamens. Histological sections show that formation of procambial vascular traces is similar in the abortive stamen primordia of [gla]tbc, the petalode primordia of `[pbg]tbc‘ and stamen primordia of male-fertile N. tabacum. The two cytoplasmic male-sterile cultivars studied here and one reported elsewhere show a common developmental stage at which the normal ontogenic pattern is altered. We suggest that this stage is the first visible point in N. tabacum androgenesis which requires genetic input from the mitochondrial genome.     

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.     

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。     

Developmental studies in orchid flowers IV: Cypripedioid species

The floral development of four species of Cypripediaceae (sensu Rasmussen 1985) was studied by means of scanning electron microscopy, with special attention to the early development of the organs that constitute the gynostemium. At the ventral base of the gynostemium a prominent structure was observed. It is most probably a vestige of the median adaxial stamen a₃ based on its early initiation and place of origin. In Cypripedium calceolus the median carpel primordium is, according to expectation, initiated slightly earlier than the lateral carpel primordia, and later develops into the largest stigma lobe. Interestingly, Cypripedium irapeanurn shows an opposite sequence in the initial phase of the carpel development in that the primordia of the lateral carpels are initiated before the primordium of the median carpel.