台闽苣苔(苦苣苔科)花部器官的形态发生

在扫描电镜下对台闽苣苔 (T. oldhamii (Hemsl.) Solereder)进行了花部器官形态发生的观察,为探索该类群的个体发育、类群间的系统发育关系和进化趋势提供依据.研究发现该属植物萼片、花冠和雄蕊发生式样均为五数花类型,它们各自来源于花原基上分化出来的萼片原基、花冠原基和雄蕊原基;花冠与雄蕊的两侧对称性与花冠上唇生长稍快和退化雄蕊原基发育迟滞相关;萼片原基的发生和发育的顺序是不一致的:萼片原基发生的式样为近轴中原基-远轴2原基-2侧原基,发育式样则为近轴中萼片-2侧萼片-远轴2萼片,花蕾时为镊合状排列.花冠裂片原基的发生和发育式样是一致的,即远轴中裂原基(下唇中裂片)-远轴2侧裂原基(下唇2侧裂片)-近轴2裂原基(上唇2裂片).花蕾期卷迭式为覆瓦状排列,从外向内:下唇中裂片-下唇2侧裂片-上唇2裂片或下唇2侧裂片-上唇2裂片-下唇中裂片.雄蕊原基与花冠裂片原基互生,前方雄蕊原基在发生上稍迟于后方雄蕊原基,后者与退化雄蕊原基几乎同时发生,但较小,并与近轴心皮(或柱头上唇)对生.将该属与玄参科(Scrophulariaceae)的地黄属( Rehmannia )、苦苣苔科(Gesneriaceae)的异叶苣苔属( Whytockia)和尖舌苣苔属(Rhynchoglossum )的花部器官比较发现,这四个属在这方面呈现出多样性和交叉.过去一直按子房室数和胎座类型划分玄参科(子房2室、中轴胎座)和苦苣苔科(子房1室、侧膜胎座)这一做法受到了质疑.     

中国广西苦苣苔科一新种--灵川小花苣苔

报道了在广西岩溶洞穴发现的苦苣苔科Gesnenaceae小花苣苔属Chiritopsis一新种——灵川小花苣苔C．lingchuanensis Yah Liu＆Y.G.Wei。本种因聚伞花序花稀疏,花冠白色,明显二唇形,上唇比下唇短3倍以上,退化雄蕊3而与羽裂小花苣苔C.bipinnatifida W .T．Wang相近,区别在于前者叶不裂,雄蕊被短柔毛。本种在叶形上还与小花苣苔C.repandaw T.Wang相近,不同在于后者花冠不明显二唇形,上下唇近等长,雄蕊无毛,退化雄蕊2。灵川小花苣苔的分布范围狭窄,目前仅见于一个岩溶洞穴的近洞口段,具明显的洞生习性。     

广西苦苣苔科稀有珍贵植物——弥勒苣苔

首次报道广西苦苣苔科植物一新记录属:弥勒苣苔属.该属接近金盏苣苔属,但不同在于弥勒苣苔属花冠上唇4浅裂,下唇不分裂,2对雄蕊分别着生于花冠中部及其上方,雌蕊具一个柱头.弥勒苣苔属为中国特有的单型属,仅弥勒苣苔一种,分布于云南东南部和广西西部.该种在广西首次记录,凭证标本存放于广西植物标本馆(IBK).     

广西苦苣苔科一新属——文采苣苔属

描述了在广西发现的苦苣苔科一新属和一新种,即文采苣苔属Wentsaiboea D. Fang & D. H. Qin及文采苣苔W. renifolia D. Fang & D. H. Qin, 并提供墨线图。文采苣苔属的柱头外形略似长檐苣苔属Dolicholoma D. Fang & W. T. Wang, 不同在于前者叶肾形,基部心形,具掌状脉,花冠斜钟状,裂片圆形,雄蕊和退化雄蕊着生于冠筒近基部。新属在体态上还接近小花苣苔属Chiritopsis W. T. Wang, 但前者叶具掌状脉,冠筒钟状,远轴侧膨胀,柱头马蹄形;在后者叶具羽状脉,冠筒筒状,不膨胀,柱头下唇倒梯形至线形。     

中国喜鹊苣苔属(苦苣苔科)一新记录种——雷氏喜鹊苣苔

报道了中国苦苣苔科(Gesneriaceae)喜鹊苣苔属(Ornithoboea Parish ex C.B.Clarke)一新记录种——雷氏喜鹊苣苔(O.lacei Craib),该种与滇桂喜鹊苣苔(O.wildeana Craib)近似,其区别特征在于花冠下唇裂片顶端凹陷,退化雄蕊3。凭证标本存放于广西植物研究所标本馆(IBK)。     

贵州金盏苣苔属(苦苣苔科)一新种——万山金盏苣苔

报道了贵州金盏苣苔属Isometrum一新种,即万山金盏苣苔I.wanshanenseS.Z.He。该种以花序梗、花梗被褐色长柔毛至近无毛,花冠细筒状,紫色,喉部不缢缩,上唇稍长于下唇,雄蕊及雌蕊均无毛而与柔毛金盏苣苔I.villosumK.Y.Pan相近缘,但叶具柄,柄长0.4–1.4cm,叶片较小,倒披针形,长1.5–4×0.5–1.6cm,聚伞花序具1–4朵花,花萼裂片外面近无毛或近顶端疏被褐色长柔毛,花柱与子房近等长而不同。     

万山金盏苣苔

报道了贵州金盏苣苔属Isometrum一新种, 即万山金盏苣苔I. wanshanense S. Z. He。该种以花序梗、花梗被褐色长柔毛至近无毛, 花冠细筒状, 紫色, 喉部不缢缩, 上唇稍长于下唇, 雄蕊及雌蕊均无毛而与柔毛金盏苣苔I. villosum K. Y. Pan 相近缘, 但叶具柄, 柄长0.4-1.4 cm, 叶片较小, 倒披针形, 长1.5-4×0.5-1.6 cm, 聚伞花序具1-4朵花, 花萼裂片外面近无毛或近顶端疏被褐色长柔毛, 花柱与子房近等长而不同。     

圆唇苣苔属(Gyrocheilos)花柱侧偏弯折现象及其传粉适应机制

圆唇苣苔属(Gyrocheilos)是苦苣苔科的中国特有属,有5种,全部狭域分布在我国西南及广东的高海拔山区。圆唇苣苔属所有物种的花柱侧偏且花柱顶端呈90°弯折,使得柱头位于花开口的中央位置。这种独特的侧偏弯折花柱结构,说明圆唇苣苔属可能有着特殊的演化历史和适应机制。为揭示这种特殊的花柱侧偏弯折现象的发生范围、发育过程及其传粉适应机制,该研究在圆唇苣苔(Gyrocheilos chorisepalus)、折毛圆唇苣苔(G. retrotrichus)和微毛圆唇苣苔(G. microtrichus)3个物种中开展了花部综合征观察,并研究了广东大雾岭保护区内的折毛圆唇苣苔花发育过程、花部特征和繁育系统以及传粉过程。结果表明:(1)微毛圆唇苣苔只有花柱左偏弯折现象,而圆唇苣苔和折毛圆唇苣苔虽然大部分花是花柱左偏弯折,但在部分个体中出现了少量的花柱右偏弯折现象(占种群总花数的2%～3%)。(2)传粉观察发现,折毛圆唇苣苔在花蕾期即出现了花柱弯折现象,2个可育雄蕊的花药合生、位于花冠筒喉部中央位置,与侧偏花柱不存在左右镜像对称关系。(3)折毛圆唇苣苔的花粉胚珠比(P/O)为456.98±15.55,属于兼性异交繁育系统。折毛圆唇苣苔存在一定的传粉限制,自交授粉可以结实,但异交种子萌发率更高,可能存在近交衰退。(4)折毛圆唇苣苔的访花昆虫较少,访花频率较低,主要访花昆虫有隧蜂、熊蜂、食蚜蝇等; 熊蜂体型较大,访花时降落在弯折花柱和花瓣下唇,胸部侧面及下部能有效接触到柱头。(5)反射率结果显示,折毛圆唇苣苔花瓣反射波长范围集中在紫光和蓝紫光区域,花冠的反射波长范围与蜂类视觉范围一致且花冠筒外侧和花瓣下唇的反射强度最大,更容易吸引蜂类落置在花冠宽大的下唇; 圆唇苣苔属的花柱侧偏弯折现象可能来自近缘的长蒴苣苔属(Didymocarpus)的花柱下弯现象或镜像花(mirror-image flowers)。综上认为,这种侧偏弯折的花柱,可能通过提供昆虫降落平台,使得柱头位于花开口中央和花瓣下唇的上方位置,提高了柱头接触访花昆虫的概率,是适应高海拔地区低频率访花者的一种机制。     

广西半蒴苣苔属（苦苣苔科）一新种——红苞半蒴苣苔

报道了在广西发现的苦苣苔科半蒴苣苔属Hemiboea一新种,即红苞半蒴苣苔H. rubribracteata Z. Y. Li & Yan Liu。该新种叶形与贵州半蒴苣苔H. cavaleriei Lévl.相近,不同在于茎较粗且坚硬,总苞红色,萼片较长,花冠外面白色,无毛,下唇3裂至中部。     

云南东南部苦苣苔科一新种——红河短檐苣苔（英文）

描述了采自中国云南东南部红河县的苦苣苔科一新种——红河短檐苣苔(Tremacron hongheense),新种与该属模式种短檐苣苔(Tremacron forrestii Craib)最为相似,区别在于该新种叶片正面疏被长刚毛,而非较密的白色贴伏短柔毛和稀疏锈色长柔毛;花冠筒外面具白色短腺毛,而非近无毛;花冠檐部裂片顶端呈红色,且质地加厚(特别是上唇裂片),而非黄色且同质地;雄蕊长约1.6~1.8 cm,而非0.4~1.2 cm长;退化雄蕊长0.5~1.4 cm,而非0.2~0.4 cm长。     

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.     

The development of pistillate and perfect florets in Xeranthemum squarrosum (Asteraceae)

The formation of capitulum inflorescence with two different types of floret is an interesting issue in floral biology and evolution. Here we studied the inflorescence, floral ontogeny and development of the everlasting herb, Xeranthemum squarrosum, using epi‐illumination microscopy. The small vegetative apex enlarged and produced involucral bracts with helical phyllotaxy, which subtended floret primordia in the innermost whorl. Initiation of floret primordia was followed by an acropetal sequence, except for pistillate peripheral florets. The origin of receptacular bracts was unusual, as they derived from the floral primordia rather than the receptacular surface. The order of whorl initiation in both disc and pistillate flowers included corolla, androecium and finally calyx, together with the gynoecium. The inception of sepals and stamens occurred in unidirectional order starting from the abaxial side, whereas petals incepted unidirectionally from the adaxial or abaxial side. Substantial differences were observed in flower structure and the development between pistillate and perfect florets. Pistillate florets presented a zygomorphic floral primordium, tetramerous corolla and androecium and two sepal lobes. In these florets, two sepal lobes and four stamen primordia stopped growing, and the ovary developed neither an ovule nor a typical stigma. The results suggest that peripheral pistillate florets in X. squarrosum, which has a bilabiate corolla, could be considered as an intermediate state between ancestral bilabiate florets and the derived ray florets.     

马先蒿属花冠无喙类的花器官发生

对花冠无喙类密穗马先蒿(Pedicularis densispica)和大王马先蒿(P．rex)的花器官电镜扫描发现,两种不同花冠型(无齿和具齿)的马先蒿花部器官发生和发育初期十分相似,表现为明显的单轴对称。2个萼片原基首先发生于花顶的近轴侧位,然后沿花顶边缘向远轴端发育形成--马蹄形结构。密穗马先蒿在近轴中部又出现1枚萼片原基,随后马蹄形结构分化出4枚萼片,并与近轴中部的原基愈合后构成5齿萼片;而大王马先蒿的2齿萼片直接由马蹄形结构发育而成。5枚独立的花瓣原基随后发生,但发育相对滞后;除近轴中部位置1枚空缺外,4枚雄蕊原基与花瓣原基位置呈交互发生;2个心皮原基同时在拱形花顶的近轴和远轴端发生,剩余的花顶形成中间的隔膜,并与2个心皮形成中轴胎座。对马先蒿与金鱼草(Antirrhinum majus)和毛地黄(Digitalis purpurea)花器官发生和发育初期的特征进行了比较,讨论了马先蒿属花冠对称性变化的意义。     

Floral organogenesis of Chloranthus sessilifolius, with special emphasis on the morphological nature of the androecium of Chloranthus (Chloranthaceae)

 Floral organogenesis of Chloranthus sessilifolius K. F. Wu is described. The inflorescence primordium is dome-like in the beginning and then elongates, and bract primordia initiate almost decussately. Each floral primordium, arising from the axil of a bract, soon becomes a scale-like structure, with three primordia of androecial lobes originating from its abaxial part, and the gynoecial primordium in adaxial position. As the androecial lobes become more distinct, four thecae are already in differentiation, and the gynoecial primordium appears as a shallow disc. The androecial lobes do not extend their length until the thecae approach maturity and the stigma is differentiated. The androecial lobes are united at all the stages of development, and the entire androecium falls off as a unit at the end of anthesis. Based on these results, combined with published evidence from neobotany, palaeobotany and phylogenetic studies, the morphological nature of the androecium of Chloranthus is further discussed. Our studies support the viewpoint that the androecial structure of Chloranthus may have arisen by splitting of a single stamen with 2 marginal thecae. Received May 2, 2001 Accepted December 18, 2001     

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。     

A SCANNING ELECTRON MICROSCOPIC STUDY ON THE INITIATION AND DEVELOPMENT OF FLORAL ORGANS OF BRASSICA NAPUS (CV. WESTAR)

The initiation and development of the floral organs of Brassica napus L. (cv. Westar) were examined using the scanning electron microscope. After transition of the vegetative apex into an inflorescence apex, flower primordia were initiated in a helical phyllotactic pattern. The sequence of initiation of the floral organs in a flower bud was that of sepals, stamens, petals and gynoecium. Of the four sepal primordia, the abaxial was initiated first, followed by the two lateral and finally the adaxial primordium. The four long stamens were initiated simultaneously in positions alternating with the sepals. The two short stamens were initiated basipetal to and outside the long stamens, and opposite the lateral sepals. The petals arose on either side of the two short stamens and the gynoecium was produced from the remainder of the apex. During development, the sepal primordia curved sharply at the tips and tightly enclosed the other organs. Stamen primordia developed tetralobed anthers at an early stage while filament elongation occurred just prior to anthesis. A unique pattern of bulbous cells was present on the abaxial surface of the anther. Growth of petal primordia lagged relative to the other floral organs but expansion was rapid prior to anthesis. The gynoecium primordium was characterized by an invagination early in development. At maturity, there was differentiation of a papillate stigma, an elongated style and a long ovary marked externally by sutures and divided internally by a septum. Distinct patterns of cuticular thickenings were observed on the abaxial and adaxial surfaces of the petals and stamens and on the surface of the style. The patterns were less obvious on the sepals and ovary. Stomata were present on both surfaces of the mature sepals, on the style and restricted areas on the abaxial surface of the anthers and nectaries but were absent from the petals, the adaxial surface of the stamens and the ovary. No hairs were present on any of the floral organs.     

Floral ontogeny in Dipterygeae (Fabaceae) reveals new insights into one of the earliest branching tribes in papilionoid legumes

Flowers of Dipterygeae (Fabaceae, Papilionoideae) exhibit an unusual petaloid calyx. The two adaxial sepals are large and petaloid, and the three abaxial sepals form a three‐toothed lobe. The goal of this study was to elucidate the ontogenetic pathways of this peculiar calyx in light of the floral development of the three genera that comprise the tribe. Floral buds of Dipteryx alata, Pterodon pubescens and Taralea oppositifolia were analysed using scanning electron microscopy and light microscopy. The order of bracteole and sepal initiation varies among the species. The androecium is asymmetric. The carpel cleft is positioned to the right or to the left, and is opposite the adaxial antepetalous stamen. The peculiarity of the calyx becomes noticeable in the intermediate stages of floral development. It results from the differential growth of the sepal primordia, in which the abaxial and lateral primordia remain diminutive during floral development, compared with the adaxial ones that enlarge and elongate. Bracteoles, abaxial sepals, petals and anthers are appendiculate, except in T. oppositifolia, in which the appendices were not found in bracteoles or anthers. These appendices comprise secretory canals or cavities. Considering that the ontogenetic pathway for the formation of the petaloid calyx is similar and exclusive for Dipterygeae, it might be a potential synapomorphy for the group, with the presence of secretory canals in the appendices of abaxial and lateral sepals and petals. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 529–550.     

弯齿盾果草(紫草科)花序及花的形态发生

以弯齿盾果草不同发育时期的花芽为材料,在体视显微镜解剖观察的基础上使用扫描电镜对弯齿盾果草花序、花及果实的发育过程进行了观察。结果显示:(1)弯齿盾果草的花序是由最初的一个球形花序原基经过多次分裂形成的,且花序发生式样符合蝎尾状聚伞花序结构,而非通常所描述的镰状或螺状聚伞花序;花序发生过程中无单一主轴,花序轴是由侧枝连接而成,每一朵花原基有其对应的1枚苞片,下一花原基是从相邻的上一枚苞腋里发生,相邻两花原基交错互生。(2)花器官的发生是按照花萼原基、花冠原基、雄蕊原基和雌蕊原基的顺序发育,但雄蕊原基的花药部分发育速度要比花冠原基快,所以花器官的发育是按照花萼、雄蕊、花冠和雌蕊的顺序发育。(3)子房四深裂结构是由4个原基分别发育,而后相互靠拢而成。(4)小坚果表面的附属结构发生于子房发育后期,其背面的内外层突起分别是由生长较快的外部组织的边缘通过上部内缩和下部向外环状生长形成。     

Inflorescence and flower development in the Hedychieae (Zingiberaceae): Hedychium coccineum Smith

The inflorescence of Hedychium coccineum Smith is thyrse, and the primary bracts are initiated in a spiral phyllotactic pattern on the sides of the inflorescence dome. Cincinnus primordia are initiated on the flank of the inflorescence apex, in the axils of primary bracts. This primordium subsequently develops a bract and a floral primordium. Then, the floral primordium enlarges, flattens apically, and becomes rounded. Sepals are initiated sequentially from the rounded corner of the primordium ring sepal initiation, and the floral primordium continues to enlarge and produces a ring primordium. Later, this ring primordium separates three common primordia surrounding a central cavity. The adaxial common primordium is the first separation. This primordium produces the posterior petal and the fertile stamen. The remaining two common primordia separate and produce respectively a petal and a petaloid, the inner androecial member. As the flower enlarges, the cavity of the floral cup becomes a rounded–triangular apex; these apices are the sites of outer androecial primordium initiation. The abaxial outer androecial member slightly forms before the two adaxial members develop. But this primordium ceases growth soon after initiation, while the two posterior primordia continue growth to produce the lateral petaloid staminodes. During this stage, gynoecial initiates in the floral cup and continues to grow until extending beyond the labellum.     

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.