花叶芋(天南星科)的花器官发生

利用扫描电镜首次观察了天南星科花叶芋(Colocasia bicolor) 的花器官发生过程。花叶芋的肉穗花序由无花被的单性花构成, 雌花发生于花序基部, 雄花发生于花序上部, 中性花位于花序中间部位。雄花: 3 或4 个初生雄蕊原基轮状发生, 随后每个初生原基一分为二, 形成6或8个次生原基; 一部分次生原基在其后的发育过程中融合, 形成5 或7 枚雄蕊; 雄花发育过程中未见雌性结构的分化; 花药的分化先于花丝; 雄蕊合生成雄蕊柱。雌花: 合生心皮, 3或4个心皮原基轮状发生, 未见雄性结构的分化。中性花来源于雌雄花序过渡带上, 属于雄蕊原基的滞后发育以及发育成熟过程中的退化; 与彩叶芋属(Caladium)不同, 此过渡区未见畸形两性花。初生雄蕊原基二裂产生次生原基的次生现象在目前天南星科花器官发生中显得比较特殊, 同时初步探讨了次生原基的融合方式。     

千金榆花序及花器官发育

在扫描电镜下首次观察了桦木科鹅耳枥属千金榆花序和花的形态发生过程。千金榆雌花序由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化形成2个花原基和2个次级苞片;每个花原基分化出2个心皮原基,形成1个二心皮雌蕊;次级苞片远轴面发育快于近轴面,呈不均等的联合状;雌蕊基部有1层环状花被原基。雄花序为柔荑状,由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化出3个花原基分区,并分化形成3朵小花,小花无花被,位于两侧的小花分别有2枚雄蕊,位于中央的小花有4枚雄蕊,雄蕊共8枚,稀为10枚,该3朵小花为二歧聚伞状排列,其花基数应为2基数。     

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.     

榛属（桦木科）花序及花的形态发生

在扫描电镜下观察了桦木科榛属榛、毛榛和滇榛的花序和花的形态发生过程。榛属雌花序由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化形成2个花原基;每个花原基分化出2个心皮原基,形成二心皮雌蕊;雌蕊基部有2层花被原基,内层花被原基环状,外层花被发生于花原基近轴面和远轴面,近轴面和远轴面的花被不均等分化,外层花被发生早于内层花被。雄花序为柔荑状,由多个小聚伞花序螺旋状排列组成。每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化出2枚次级苞片和4。6个雄蕊原基,形成4—6枚雄蕊,每个雄蕊具4个药囊,在雄蕊原基分化形成4药囊雄蕊过程中．出现雄蕊原基纵裂。并且花丝纵裂至基部。为进一步全面探讨桦木科属间系统演化关系提供了证据。     

榛属 (桦木科) 花序及花的形态发生

在扫描电镜下观察了桦木科榛属榛、毛榛和滇榛的花序和花的形态发生过程。榛属雌花序由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化形成2个花原基;每个花原基分化出2个心皮原基,形成二心皮雌蕊;雌蕊基部有2层花被原基,内层花被原基环状,外层花被发生于花原基近轴面和远轴面,近轴面和远轴面的花被不均等分化,外层花被发生早于内层花被。雄花序为柔荑状,由多个小聚伞花序螺旋状排列组成。每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化出2枚次级苞片和4~6个雄蕊原基,形成4~6枚雄蕊,每个雄蕊具4个药囊,在雄蕊原基分化形成4药囊雄蕊过程中,出现雄蕊原基纵裂,并且花丝纵裂至基部。为进一步全面探讨桦木科属间系统演化关系提供了证据。     

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.     

Ontogeny of Staminate and Carpellate Flowers of Schisandra sphenanthera(Schisandraceae)

The ontogenetic process of the staminate and carpellate flowers of Schisandra sphenanthera Rehd. et Wils., an endemic species to China, was observed for the first time under the scanning electron microscope (SEM). In the staminate flowers, the perianth units and stamens were initiated acropetally in a continuous fasion with 2/5 spiral phyllotaxis, while no female structures were formed. Anthers were differentiated prior to the filaments formation. Throughout all the stages were the stamens arranged spirally on a columniform receptacle. In the carpellate flowers, the initiation sequence of the perianth units and carpels were similar to that of the staminate flowers. In contrast, no male structures were formed. Shortly after initiation, the carpel primordia began their marginal growth besides the apical growth and then appresses were formed on the adaxial surfaces of the primordia. However the lower margins of these appresses were inconspicuous, resulting in conduplicate carpels. Two ovules were developed on the inner surface near either lateral margins of the carpel, shaping laminar placentae. Compared with S. glabra (Brickell) Rehd., a related American species, the evolutionary trend of phyllotaxis of androecia is considered that stamens may change from spiral to approximately whorled arrangement, accompanying with the change of receptacle from a column to a flattened shield. It was also suggested that the stamens being numerous and uncertain in number become certain and decrease in number to 5 (4－7). Sterile stamens are observed and the unisexual nature of the flowers is discussed. Two types of carpel primordia are categorized, corresponding to two types of carpels, namely, ascidiate and conduplicate carpels, respectively.     

INFLORESCENCE AND FLORAL DEVELOPMENT IN HOUTTUYNIA CORDATA (SAURURACEAE)

The inflorescence of Houttuynia cordata produces 45–70 sessile bracteate flowers in acropetal succession. The inflorescence apical meristem has a mantle-core configuration and produces “common” or uncommitted primordia, each of which bifurcates to form a floral apex above, a bract primordium below. This pattern of organogenesis is similar to that in another saururaceous plant, Saururus cernuus. Exceptions to this unusual development, however, occur in H. cordata at the beginning of inflorescence activity when four to eight petaloid bract primordia are initiated before the initiation of floral apices in their axils. “Common” primordia also are lacking toward the cessation of inflorescence apical activity in H. cordata when primordia become bracts which may precede the initiation of an axillary floral apex. Many of these last-formed bracts are sterile. The inflorescence terminates with maturation of the meristem as an apical residuum. No terminal flowers or terminal gynoecia were found, although subterminal gynoecia or flowers in subterminal position may overtop the actual apex and obscure it. Individual flowers have a tricarpellate syncarpous gynoecium and three stamens adnate to the carpels; petals and sepals are lacking. The order of succession of organs is: two lateral stamens, median stamen, two lateral carpels, median carpel. The three carpel primordia almost immediately are elevated as part of a gynoecial ring by zonal growth of the receptacle below the attachment of the carpels. The same growth elevates the stamen bases so that they appear adnate to the carpels. The trimerous condition in Houttuynia is the result of paired or solitary initiations rather than trimerous whorls. Symmetry is bilateral and zygomorphic rather than radial. No evidence of spiral arrangement in the flower was found.     

桂味荔枝花器官的发生和发育过程研究

利用SV11立体显微镜和JSM-6360LV型扫描电镜观察‘桂味’荔枝花器官的发生和发育过程。结果表明:花序原基最先发生,然后形成数个大小不等的单花原基;4个萼片原基的发生不同步,其中一侧对位先发生;6～10枚雄蕊原基以轮状方式几乎同时发生;心皮原基最后发生,2～3枚(稀4枚)心皮原基同时出现,随后进行侧向生长,逐渐合拢形成子房。雌花中,花柱、柱头分化明显,雄蕊退化。雄花中,花丝细长,花药饱满,雌蕊退化或发育不完全。两性花中,雌雄蕊发育完全。花粉粒近球形,具3孔沟,表面为条纹状纹饰。     

Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb

The annual Mediterranean herbTheligonum cynocrambe shows a peculiar combination of morphological characters, e.g., switch from decussate to spiral phyllotaxis with 90–100° divergence, combined with a change from interpetiolar to lateral stipules, anemophily, lack of calyx, flowers often dimerous to trimerous, corolla fused in both male and female flowers, male flowers extra-axillary, with 2–19 stamens per flower, female flowers axillary, with inferior uniovulate ovary, basilateral style and perianth, nut-like fruits with elaiosome. In male flowers the androecium emerges as an (uneven) elliptical rim with a central depression. This common girdling primordium is divided up into several stamen primordia. In male flowers with low stamen number the stamen primordia may occupy the corners alternating with the corolla lobes. There are no epipetalous androecial primordia that secondarily divide into stamens. Male flowers occasionally show a hemispherical base that may be interpreted as remnant of the inferior ovary. In female flowers a ring primordium grows into a tube on which the petal lobes arise. The perianth and style become displaced adaxially by uneven growth of the inferior ovary. The ovary is basically bilocular. The lower region of the ovary is provided with a septum that is overtopped and hidden by the single curved ovule.Theligonum is referred to theRubiaceae-Rubioideae, with theAnthospermeae andPaederieae as most closely related tribes.     

The need to re-investigate the nature of homoplastic characters: an ontogenetic case study of the 'bracteoles' in Atripliceae (Chenopodiaceae)

Background and Aims

Within Chenopodioideae, Atripliceae have been distinguished by two bracteoles enveloping the female flowers/fruits, whereas in other tribes flowers are described as ebracteolate with persistent perianth. Molecular phylogenetic hypotheses suggest ‘bracteoles’ to be homoplastic. The origin of the bracteoles was explained by successive inflorescence reductions. Flower reduction was used to explain sex determination. Therefore, floral ontogeny was studied to evaluate the nature of the bracteoles and sex determination in Atripliceae.

Methods

Inflorescences of species of Atriplex, Chenopodium, Dysphania and Spinacia oleracea were investigated using light microscopy and scanning electron microscopy.

Key Results

The main axis of the inflorescence is indeterminate with elementary dichasia as lateral units. Flowers develop centripetally, with first the formation of a perianth primordium either from a ring primordium or from five individual tepal primordia fusing post-genitally. Subsequently, five stamen primordia originate, followed by the formation of an annular ovary primordium surrounding a central single ovule. Flowers are either initially hermaphroditic remaining bisexual and/or becoming functionally unisexual at later stages, or initially unisexual. In the studied species of Atriplex, female flowers are strictly female, except in A. hortensis. In Spinacia, female and male flowers are unisexual at all developmental stages. Female flowers of Atriplex and Spinacia are protected by two accrescent fused tepal lobes, whereas the other perianth members are absent.

Conclusions

In Atriplex and Spinacia modified structures around female flowers are not bracteoles, but two opposite accrescent tepal lobes, parts of a perianth persistent on the fruit. Flowers can achieve sexuality through many different combinations; they are initially hermaphroditic, subsequently developing into bisexual or functionally unisexual flowers, with the exception of Spinacia and strictly female flowers in Atriplex, which are unisexual from the earliest developmental stages. There may be a relationship between the formation of an annular perianth primordium and flexibility in floral sex determination.     

THE ONTOGENY OF THE INFLORESCENCE AND FLOWER OF LIQUIDAMBAR STYRACIFLUA L. (HAMAMELIDACEAE)

A developmental study of the inflorescence of Liquidambar styraciflua L. was conducted to clarify morphological discrepancies reported in the literature. Salient features of development are: 1) the inflorescence apex results from the conversion of a terminal, vegetative apex; 2) partial inflorescence apices arise as ellipsoid structures in axils of leaves, bracts, or transitional phyllomes; 3) development of male heads is acropetal whereas female heads differentiate basipetally; 4) the partial inflorescence apex becomes segmented into several distinct subunits indicating an axillary branch system of the third order; 5) distinct individual floral primordia are initiated on the subunits; 6) a complete absence of perianth development; 7) inception of carpel primordia in flowers of lower male heads as well as female heads, but a failure of the gynoecium to develop beyond an incipient stage in male heads; and 8) development of sterile structures around the base of the styles of only female flowers near the time of anthesis. Carpellary characteristics of the sterile structures are described, their morphological nature is discussed, and the phylogenetic position of Liquidambar is evaluated.     

Expression of ASK1-like genes in arrested stamens of female Silene latifolia plants

Silene latifolia is a dioecious plant in which sex is determined by heteromorphic sex chromosomes. In female plants, stamen development is arrested before microspore mother cells are formed. In this study, we isolated four cDNAs (SlSKP1-1 to 4) encoding ASK1-like protein as expression markers to reveal when expression levels are reduced in arrested stamens of female flowers. Expression patterns of the SlSKP1 genes were analyzed by in-situ hybridization. We use the flower development classification of Grant et al. (in Plant J 6:471–480, 1994). SlSKP1 genes were highly expressed in primary parietal cells and primary sporogenous cells that develop into microspore mother cells in male flowers. Expression levels started to be reduced in the external stamens of the female flowers when stamen development was arrested at stage 7. Although microspore mother cells could not be developed in female flowers and SlSKP1 expression may be unnecessary in arrested stamens, SlSKP1 genes were still expressed in sporogenous cells of degenerated stamens at stage 8. Parietal cells stopped differentiating earlier than sporogenous cells in arrested stamens. These results suggest that not all types of cell are arrested simultaneously at a particular stage of stamen development during stamen suppression of S. latifolia.     

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.     

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.     

INITIATION AND DEVELOPMENT OF INFLORESCENCE AND FLOWER IN ANEMOPSIS CALIFORNICA (SAURURACEAE)

All flowers of Anemopsis californica, the most specialized taxon of the family Saururaceae, are initiated as individual primordia subtended by previously initiated bracts, in contrast to the common-primordium initiation of all flowers of Saururus cernuus and of most flowers of Houttuynia cordata. Floral symmetry is bilateral and zygomorphic, and the sequence of initiation among floral parts is paired or whorled. In A. californica, the six stamens arise as three common primordia, each of which later bifurcates to form a pair. The three common primordia occupy sites corresponding to the positions of the three stamens in H. cordata flowers. In Anemopsis, the filaments of each pair are connate. Each stamen pair is vascularized by a single bifurcating vascular bundle. The three carpels per flower are usually initiated simultaneously although there may be some variation. Adnation between stamens and carpels results from zonal growth. Downward extension of the locule, and proliferation and expansion of receptacular tissue and inflorescence cortical tissue around the locule below the bases of the carpels produce the inferior ovary. The inflorescence terminates its activity as a flattened apical residuum, surrounded by bracts subtending reduced flowers most of which have stamens only.     

铁木属(桦木科)花序及花的形态发生

在扫描电镜下观察了桦木科(Betulaceae)铁木属花序和花的形态发生过程。结果显示, 铁木雌花序由多个小聚伞花序螺旋状排列组成。每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织, 由小花序原基分生组织分化形成1对次级苞片和2个花原基, 每个花原基分化出2个或3个心皮原基, 形成二心皮或三心皮雌蕊, 雌蕊基部有1层环状花被原基。雄花序为柔荑状, 由多个小聚伞花序螺旋状排列组成。每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织, 由小花序原基分生组织分化出3个花原基分区, 位于中央的花原基分区, 分化形成5-6枚雄蕊原基, 两侧的花原基分区, 分别分化形成3-4枚雄蕊原基, 雄蕊原基分化形成四药囊雄蕊。雄蕊原基纵裂, 但花丝纵裂没有达到基部。     

Floral ontogeny and systematic position of the Didiereaceae

Floral ontogenetical data from all four genera of the Didiereaceae (s.str.) are presented for the first time. All Didiereaceae s.str. are dioecious, having unisexual flowers with organ rudiments of the opposite sex. Two median bracts followed by a tetramerous perianth (two alternating dimerous ``whorls'), a slightly complex androecium with 6–12 stamens in a single row (on a common ring primordium), four of which mostly alternating with the perianth members, and one basal ovule connecting three free septa at their very base are flower characters in Didiereaceae, supporting phylogenetic analyses based on nucleotide sequence data. Closest relatives are the (formerly) portulacaceous genera Portulacaria (5 stamens alternating with the perianth), Ceraria (5 stamens alternating with the perianth), and Calyptrotheca (many stamens), all with pentamerous perianths, from which the tetramerous perianth in Didiereaceae can be derived. Applequist and Wallace (2003) included these three genera in an expanded family Didiereaceae (with three subfamilies).     

平基槭花性别分化的细胞形态学观察

平基槭为杂性花,雄花与两性花同株,本文对其花性别分化过程进行了细胞形态学观察。结果发现,在花性别分化的早期,雄花和两性花的花芽中雌、雄蕊原基均具备,只是在花芽发育到一定时期,雄花的雌蕊原基发生选择性败育,败育发生在大孢子母细胞减数分裂为4个大孢子时期。两性花的雌蕊可以正常膨大结实,雄蕊花药虽然可以形成二核花粉,但不能正常开裂,属于不育雄蕊。初步分析认为,两性花雄蕊花药不能正常开裂与花粉囊壁纤维层木质化程度低有关。     

Comparative floral development in male and female plants of Palmer amaranth (Amaranthus palmeri)

Premise

Characterizing the developmental processes in the transition from hermaphroditism to unisexuality is crucial for understanding floral evolution. Amaranthus palmeri, one of the most devastating weeds in the United States, is an emerging model system for studying a dioecious breeding system and understanding the biological traits of this invasive weed. The objectives of this study were to characterize phases of flower development in A. palmeri and compare organogenesis of flower development in female and male plants.

Methods

Flower buds from male and female plants were dissected for light microscopy. Segments of male and female inflorescences at different stages of development were cut longitudinally and visualized using scanning electron microscopy.

Results

Pistillate flowers have two to three styles, one ovary with one ovule, and five obtuse tepals. Staminate flowers have five stamens with five acute tepals. Floral development was classified into 10 stages. The distinction between the two flower types became apparent at stage four by the formation of stamen primordia in staminate flowers, which developed female and male reproductive organs initially, as contrasted to pistillate flowers, which produced carpel primordia only. In staminate flowers, the putative carpel primordia changed little in size and remained undeveloped.

Conclusions

Timing of inappropriate organ termination varies across the two sexes in A. palmeri. Our study suggests that the evolution of A. palmeri from a cosexual ancestral state to complete dioecy is still in progress since males exhibited transient hermaphroditism and females produced strictly pistillate flowers.