Floral development and vasculature in Eriocaulon (Eriocaulaceae) provide insights into the evolution of Poales

Background and AimsFloral developmental studies are crucial for understanding the evolution of floral structures and sexual systems in angiosperms. Within the monocot order Poales, both subfamilies of Eriocaulaceae have unisexual flowers bearing unusual nectaries. Few previous studies have investigated floral development in subfamily Eriocauloideae, which includes the large, diverse and widespread genus Eriocaulon. To understand floral variation and the evolution of the androecium, gynoecium and floral nectaries of Eriocaulaceae, we analysed floral development and vasculature in Eriocaulon and compared it with that of subfamily Paepalanthoideae and the related family Xyridaceae in a phylogenetic context.MethodsThirteen species of Eriocaulon were studied. Developmental analysis was carried out using scanning electron microscopy, and vasculature analysis was carried out using light microscopy. Fresh material was also analysed using scanning electron microscopy with a cryo function. Character evolution was reconstructed over well-resolved phylogenies.Key ResultsPerianth reductions can occur due to delayed development that can also result in loss of the vascular bundles of the median sepals. Nectariferous petal glands cease development and remain vestigial in some species. In staminate flowers, the inner stamens can emerge before the outer ones, and carpels are transformed into nectariferous carpellodes. In pistillate flowers, stamens are reduced to staminodes and the gynoecium has dorsal stigmas.ConclusionsFloral morphology is highly diverse in Eriocaulon, as a result of fusion, reduction or loss of perianth parts. The nectariferous carpellodes of staminate flowers originated first in the ancestor of Eriocaulaceae; petal glands and nectariferous branches of pistillate flowers originated independently in Eriocaulaceae through transfer of function. We present a hypothesis of floral evolution for the family, illustrating a shift from bisexuality to unisexuality and the evolution of nectaries in a complex monocot family, which can contribute to future studies on reproductive biology and floral evolution in other groups.     

Pre-adaptations and the evolution of pollination by sexual deception: Cope's rule of specialization revisited

Pollination by sexual deception is arguably one of the most unusual liaisons linking plants and insects, and perhaps the most illustrative example of extreme floral specialization in angiosperms. While considerable progress has been made in understanding the floral traits involved in sexual deception, less is known about how this remarkable mimicry system might have arisen, the role of pre-adaptations in promoting its evolution and its extent as a pollination mechanism outside the few groups of plants (primarily orchids) where it has been described to date. In the Euro-Mediterranean region, pollination by sexual deception is traditionally considered to be the hallmark of the orchid genus Ophrys. Here, we introduce two new cases outside of Ophrys, in plant groups dominated by generalized, shelter-mimicking species. On the basis of phylogenetic reconstructions of ancestral pollination strategies, we provide evidence for independent and bidirectional evolutionary transitions between generalized (shelter mimicry) and specialized (sexual deception) pollination strategies in three groups of flowering plants, and suggest that pseudocopulation has evolved from pre-adaptations (floral colours, shapes and odour bouquets) that selectively attract male pollinators through shelter mimicry. These findings, along with comparative analyses of floral traits (colours and scents), shed light on particular phenotypic changes that might have fuelled the parallel evolution of these extraordinary pollination strategies. Collectively, our results provide the first substantive insights into how pollination sexual deception might have evolved in the Euro-Mediterranean region, and demonstrate that even the most extreme cases of pollinator specialization can reverse to more generalized interactions, breaking ‘Cope''s rule of specialization’.     

Phytochromes A1 and B1 have distinct functions in the photoperiodic control of flowering in the obligate long-day plant Nicotiana sylvestris

The obligate long-day plant Nicotiana sylvestris with a nominal critical day length of 12 h was used to dissect the roles of two major phytochromes (phyA1 and phyB1) in the photoperiodic control of flowering using transgenic plants under-expressing PHYA1 (SUA2), over-expressing PHYB1 (SOB36), or cosuppressing the PHYB1 gene (SCB35). When tungsten filament lamps were used to extend an 8 h main photoperiod, SCB35 and SOB36 flowered earlier and later, respectively, than wild-type plants, while flowering was greatly delayed in SUA2. These results are consistent with those obtained with other long-day plants in that phyB has a negative role in the control of flowering, while phyA is required for sensing day-length extensions. However, evidence was obtained for a positive role for PHYB1 in the control of flowering. Firstly, transgenic plants under-expressing both PHYA1 and PHYB1 exhibited extreme insensitivity to day-length extensions. Secondly, flowering in SCB35 was completely repressed under 8 h extensions with far-red-deficient light from fluorescent lamps. This indicates that the dual requirement for both far-red and red for maximum floral induction is mediated by an interaction between phyA1 and phyB1. In addition, a diurnal periodicity to the sensitivity of both negative and positive light signals was observed. This is consistent with existing models in which photoperiodic time measurement is not based on the actual measurement of the duration of either the light or dark period, but rather the coincidence of endogenous rhythms of sensitivity - both positive and negative - and the presence of light cues.     

Ioral organogenesis of Echinodorus nazonicus Rataj and floral construction f the Alismatales

The primary trimerous pattern of the flower of Echinodorus amazonicus Rataj is clearly exhibited in the earliest developmental stages. After the inception of three sepals, theree alternisepalous petal-stamen complexes arise, each of which, at successively higher levels of the primordia, consists of a petal, a pair of stamens, and a single stamen. In alternation with these three petal-stamen complexes three groups of three pistil (carpel) primordia are formed. Subsequent pistil primordia are formed rapidly in alternation with the preceding ones. Teratological buds were observed in which one pistil primordium appeared to be replaced by a normal stamen primordium. General features of the flowers of the Alismatales are summarized in developmental fashion which includes the mature flower as a final stage. With the exception of Limnocharis and probably Wisneria , all alismatalean flowers appear to be characterized by antipetalous stamen (or staminodial) paris. In members studied thus far, the basic trimery differs radically from that found in certain taxa of the Magnoliidae.     

The occurrence and structure of apparently bisexual flowers in the date palm, Phoenix dactylifera L. (Arecaceae)

Individuals of Phoenix dactylifera L. have expanded pistillodes or pseudocarpels in staminate flowers. These pseudocarpels are located in the centre of the male flowers and are surrounded by stamens. The gynoecium has the characteristic three carpellate arrangement commonly found in female date palm flowers. Pseudocarpels from male flower buds can expand into parthenocarpic fruit. Histology of the expanded pistillodes or pseudotarpels is similar to that of normal carpels from pistillate plants. These pseudocarpels lack ovules. Nutrient medium containing 10 mg 1^-1 of 2,4-dichlorophenoxyacetic acid or p-chlorophenylacetic acid and 0.3% activated neutralized charcoal enhanced the development and outgrowth of the pseudocarpels of cultured male flowers.     

花颜色和花气味的量化研究方法

花颜色和花气味是花部构成的重要内容。在已开展的传粉生态学研究中对二者的报道主要是描述性的,而其量化研究可以为揭示传粉机制提供有力的实验证据。本文主要介绍了花颜色的测量和标定方法,包括比色卡、分光色差仪和便携式光谱仪等;花气味的采集方法,包括动态顶空套袋-吸附采集法、吸附-溶剂洗脱法和固相微萃取法等;花气味的检测和分析方法,包括气相色谱-质谱联用仪分析和电子鼻型超速气相色谱仪分析等;以及昆虫行为学实验方法,包括气相色谱-昆虫触角电位联用技术、Y型嗅觉仪和飞行箱实验等。科研人员可以根据实验材料的特点和实验目的选择适合的量化研究方法。     

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。     

天南星科植物的花多样性与传粉策略

天南星科植物具有特殊的佛焰苞花序及多样化的传粉策略, 是研究被子植物花的分化与动植物之间进化生态学联系的理想材料。本文简述了天南星科不同类型的花序结构及其传粉适应意义, 总结了天南星科传粉策略的基本类型与演化历史。天南星科的苞片结构主要包括原始型、外展平面型、直立宽佛焰苞型和直立狭佛焰苞4种类型, 呈现出从简单的片状与外展平面状结构向复杂的立体包裹状的佛焰苞结构演化的趋势。肉穗花序可分为两性花花序、单性花雌雄同序和单性花雌雄异序3种类型, 演化路线为两性花花序→单性花雌雄同序→单性花雌雄异序。天南星科的传粉者主要有鞘翅目、双翅目、膜翅目昆虫, 表现出5种主要传粉策略: 食物报酬型互利传粉、气味吸引型欺骗性传粉、交配场所型互利传粉、产卵场所型互利传粉和致死陷阱型欺骗性传粉。天南星科植物通过花序的形状、颜色、产热以及花部挥发物来吸引传粉者, 其中最主要的挥发物有二甲基硫化物、甲基吲哚化合物、萜类和苯类化合物, 模拟食物或产卵场所信号吸引鞘翅目甲虫和双翅目昆虫为其传粉。天南星科植物的佛焰苞被认为是促进该科物种分化的一个重要结构, 但该性状的演化历史及其与传粉系统分化之间的内在联系尚不明确。利用现代分子生物学技术以及模型模拟等手段, 结合生理生态学方法深入探究传粉事件与天南星科植物的花多样性以及物种分化之间的联系, 有望提升关于植物-传粉者互作与植物的花多样性分化之间关系的认识, 并丰富对被子植物多样性演化相关研究的理解。     

Models of floral pattern in detached flowers of Silene coeli-rosa (L) Godr. (Caryophyllaceae)

Organ number per whorl was analysed in aberrant flowers of the long-day (LD) plant , Silene coeli-rosa , to test a hypothesis that organ number in a whorl takes its cue from an adjacent outer whorl and that perturbed organ number per whorl is not random but defaults to that of closely related taxa or genera of the Caryophyllaceae. When plants were grown under short-days (SD), transferred to LD and the shoot meristem excised and cultured in vitro under SD, the normal pattern of flower development was often disrupted. For example, we observed flowers which comprised floral whorls with an aberrant number of floral organs. In part, this was an effect of tissue culture; however, the over-and-above effect was the establishment of an alternative pattern of development. Our data indicate that two distinct and recurrent patterns occurred in the aberrant flowers we observed in five separate experiments. First, pairs of floral whorls were linked so that aberration in one whorl resulted in the next whorl being more aberrant than normal. Second, the number of organs in aberrant whorls was not random, but defaulted to an organ number which mimicked the flowers of closely related species of Silene or related genera in the Caryophyllaceae.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 140 , 229−235.     

Floral morphogenesis of the Maddenia and Pygeum groups of Prunus (Rosaceae), with an emphasis on the perianth

Although the vast majority of Prunus L. (Rosaceae) species have clearly differentiated sepals and petals, two former genera Maddenia and Pygeum have been described as having an undifferentiated perianth. However, floral morphological and morphogenetic data are scarce, and a renewed investigation is essential to understand the evolution of the perianth differentiation. Here, floral morphogenesis in Prunus hypoleuca (Koehne) J.Wen (=Maddenia hypoleuca Koehne) and Prunus topengii (Merr.) J. Wen & L. Zhao (=Pygeum topengii Merr.) were examined with scanning electron microscopy. The floral development demonstrates that the ten perianth parts can be distinguished as five sepals in an external whorl and five petals in an internal whorl. The sepal primordia are broad, crescent-shaped, and truncate. The petal primordia are rounded and initially resemble the androecium. However, at maturity petals and sepals look much the same in the two species, differing from other Prunus species. The ovule is anatropous and unitegmic, but there is a basal appendage near the ovule of P. hypoleuca which is absent in P. topengii. The direction of development of floral nectaries in the hypanthium is basipetal in P. hypoleuca but acropetal in P. topengii. Perianth segments are differentiated in the two groups and the similarity of the perianth parts is secondarily acquired. Our results support the separation of the Maddenia and Pygeum groups as well as their inclusion in a broader monophyletic Prunus based on molecular phylogenetic studies. We herein provide a new nomenclatural change: Prunus topengii (Merr.) J. Wen & L. Zhao, comb. nov.     

Chaotic Floral Phyllotaxis and Reduced Perianth in Achlys (Berberidaceae)

Among the 16 genera of the Berberidaceae Achlys is the only one with a reduced perianth, an irregular floral phyllotaxis, and variable stamen number. Early floral stages show an unstable (chaotic) arrangement of the organ primordia. Only the single carpel of the gynoecium has a more fixed position in that the placenta is formed in the adaxial half of the flower. The irregularities in the androecium may be caused by the lack of influence of a perianth on floral symmetry. On the other hand, the regular orientation of the carpel is perhaps due to the early polarity of the flower, whereby the abaxial half of the flower is larger (with further developed stamen primordia) at the time when carpel polarity is established.     

APETALA1启动子驱动AtIPT4在转基因拟南芥中表达导致花和花器官发育异常

细胞分裂素对拟南芥（Arabidopsis thaliana）花分生组织细胞的分裂和分化具有重要作用。本研究利用APETALA1（AP1）特异启动子在花分生组织和第1、2轮花器官中表达细胞分裂素合成酶（isopentyl transferase,IPT）基因IPT4,研究细胞分裂素对花和花器官发育的影响。在pAP1∷IPT4转基因植株中出现了花密集和花器官数目增多等现象。原位杂交和GUS组织染色结果发现,在pAP1∷IPT4转基因植株中,花分生组织特征决定基因LEAFY（LFY）与花器官特征决定基因AP1、PISTILLATA（PI）和AGAMOUS（AG）的表达量均有不同程度的提高。研究结果表明在拟南芥中表达pAP1∷IPT4影响其花和花器官的正常发育。     

Partial floral evocation by high irradiance in the long-day plant Sinapis alba

Previous work has shown that increased irradiance promotes flower initiation in the long-day plant Sinapis alba when given during the first half of a single, suboptimal long day. It is without flower-promoting effect, however, when given during a single short day. In the present work it is shown that, in the apex of plants exposed to a single short day at high irradiance, some changes occur that are normally observed during the transition to flowering (full evocation), e.g., elevated soluble sugar and starch levels, increased numbers of mitochondria and changed nucleolus structure. These changes are of similar magnitude and follow the same sequence as the corresponding changes during full evocation. Other changes, normally associated with full evocation, e.g., increased mitotic activity, are not caused by one short day at high irradiance. This treatment thus produces only what we call "partial evocation".     

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.     

Experimental control of floral reversion in isolated shoot apices of the long-day plant Silene coeli-rosa

The fate of shoot meristems of the long day (LD) plant Silene coeli-rosa in culture was examined (complete, reverted or arrested flowers) to establish whether these different patterns were related to a particular stage of morphogenesis and the extent to which the fate of the pattern was regulated by either added plant growth regulators (PGRs) or changing the carbohydrate source in the medium. In particular, the frequency of reversion was measured to test the stability of the determined state for each whorl. The plants were given various inductive treatments (4–7 LD, 7 LD + 1 to 3 SD) and the apices were explained onto Murashige-Skoog medium supplemented with 3% sucrose (controls) ± IAA, ± kinetin, ± GA₃ or onto the basal medium containing 1 or 3% sucrose, glucose or fructose or 7% sucrose. The apices were examined 12 weeks later. When the data were pooled from all inductive treatments, IAA resulted in more reversions, GA₃ caused more arrested flowers while kinetin hardly affected the pattern of meristem fate compared with the controls. However, each PGR treatment did not perturb the pattern of organ formation for those apices that formed either arrested or complete flowers. The time for determination (days) of the earlier formed whorls (determination times for the controls in brackets): sepals (2), stamens 1–5 (3) and petals (3), was shortened by about a day in all PGR treatments whereas the corresponding times for the later formed whorls: stamens 6–10 (4) and carpels (4), were either lengthened to 5 days or unaffected. The response of the apices to the various sugars was simply a reflection of concentration. Hence, more complete flowers formed at 7 or 3% and more flowers were arrested at 1 % regardless of the sugar moiety. However, the frequency of reversion was similar on each of the media. Pooling all data from all treatments enabled a statistical analysis of the pattern of reversion and the pattern of arrest. Reversion was more common from apices which exhibited the later-formed whorls (stamens 6–10 and carpels) than from the earlier whorls. Moreover, the stronger the inductive treatment the less frequent was reversion. The most common stage of arrest was at the stamen 6–10 whorl and this was particularly so for the GA₃ treatment. The data indicated that reversion could occur from any whorl, which suggests that determination of each whorl is independent of the next. This conclusion is underlined by the more frequent occurrence of reversion from the carpel whorl. However, the longer the inductive treatment the less likelihood of reversion; this suggests that in Silene, the floral stimulus is required continuously to stabilise the determined state of each whorl and to ensure smooth completion of floral morphogenesis.     

Floral organogenesis of Helleborus thibetanus and Nigella damascena (Ranunculaceae) and its systematic significance

Floral organogenesis in Helleborus thibetanus and Nigella damascena was examined and compared using scanning electron microscopy and light microscopy, and the putative relationships of Helleborus and Nigella were analysed. H. thibetanus and N. damascena share some features of floral phyllotaxis and development of the sepals, petals, stamens and carpels, which are also found in other members of Ranunculaceae. However, they differ strongly in the number and degree of fusion of the carpels: in H. thibetanus, the two carpels are slightly united at the base, whereas, in N. damascena, the gynoecium is syncarpous and the five carpels are united throughout the ovary. Differences are also noted in petal development. The blade of the young petal of H. thibetanus develops two bulges which become connate and then fuse with the blade at the sides, developing more quickly than the blade and forming a tubular petal. In N. damascena, a single ridge is formed on the petal blade which develops into the smaller adaxial labium of the bilabiate petal, whereas the blade itself develops into the larger abaxial labium bearing two pseudonectaries. The outermost stamens are delayed in development in Helleborus, but not in Nigella. Although the results from our investigation are preliminary, differences in floral development characters suggest that Helleborus and Nigella may not be closely related and possibly support placement into separate tribes. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 431–443.     

The adaptedness of the floral phenotype in a relict endemic, hawkmoth-pollinated violet. 1. Reproductive correlates of floral variation

This paper examines the relationship between quantitative variation in floral morphology (sizes of petals, spur and peduncle) and maternal reproductive success (seed production) in Viola cazorlensic (Violaceae), a narrowly endemic violet of south-eastern Spain pollinated by day-flying hawkmoths (Sphingidae). This plant is characterized by broad intraspecific variation in size and proportions of floral parts. Floral morphology does not influence significantly the probability of fruit set. Among flowers setting fruit, spur length and size of petals have no significant effect on seed production, but capsules from long- and short-peduncted flowers contain significantly more seeds than capsules from flowers with intermediate peduncles. Individual plants differ significantly in average floral characteristics. Plants with comparatively long and short peduncles tend to produce more seeds than those with intermediate ones, even after accounting statistically for individual differences in flower production. These findings are interpreted as evidence of disruptive selection on peduncle length during the study season. Floral variability in this species may be explained by the combined action of disruptive selection on peduncle length (the character most variable among individuals) and little, if any, stabilizing selection on spur length and size of petals