Ontogeny of the proliferous spikelet in Eleocharis viridans (Cyperaceae)

Cyperaceae, the third largest family of monocotyledons, show unusual asexual reproductive strategies, such as pseudovivipary involving the formation of new individuals from somatic tissues of floral structures. In Eleocharis pseudovivipary is an important reproductive process in series Tenuissimae, which results in a structure called a proliferous spikelet. With the aim of describing the ontogenetic process of the pseudovivipary in E. viridans, proliferous spikelets at different developmental stages were analysed by light microscopy and scanning electron microscopy. The proliferous spikelet develops from a meristem located in the axil of the basal glume, homologous to bracts of other groups of Cyperaceae, in the culm of the parental plant. Each proliferous spikelet is formed by sympodial units, which consist of an addorsed prophyll, an outer and inner sheath and a culm, which develops a primordium of floriferous spikelet at the terminal region, which can be aborted, and may develop a proliferous spikelet lateral in the axil of the basal glume. The second internode of each sympodial unit contains a root primordium and an intercalary meristem at the culm base. Our results indicate that pseudovivipary can coexist with sexual reproduction as an alternative reproductive strategy, allowing the rapid spread of populations. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 524–539.     

Spikelet structure and development in Cyperoideae (Cyperaceae): a monopodial general model based on ontogenetic evidence

Background and Aims

In Cyperoideae, one of the two subfamilies in Cyperaceae, unresolved homology questions about spikelets remained. This was particularly the case in taxa with distichously organized spikelets and in Cariceae, a tribe with complex compound inflorescences comprising male (co)florescences and deciduous female single-flowered lateral spikelets. Using ontogenetic techniques, a wide range of taxa were investigated, including some controversial ones, in order to find morphological arguments to understand the nature of the spikelet in Cyperoideae. This paper presents a review of both new ontogenetic data and current knowledge, discussing a cyperoid, general, monopodial spikelet model.

Methods

Scanning electron microscopy and light microscopy were used to examine spikelets of 106 species from 33 cyperoid genera.

Results

Ontogenetic data presented allow a consistent cyperoid spikelet model to be defined. Scanning and light microscopic images in controversial taxa such as Schoenus nigricans, Cariceae and Cypereae are interpreted accordingly.

Conclusions

Spikelets in all species studied consist of an indeterminate rachilla, and one to many spirally to distichously arranged glumes, each subtending a flower or empty. Lateral spikelets are subtended by a bract and have a spikelet prophyll. In distichously organized spikelets, combined concaulescence of the flowers and epicaulescence (a newly defined metatopic displacement) of the glumes has caused interpretational controversy in the past. In Cariceae, the male (co)florescences are terminal spikelets. Female single-flowered spikelets are positioned proximally on the rachis. To explain both this and the secondary spikelets in some Cypereae, the existence of an ontogenetic switch determining the development of a primordium into flower, or lateral axis is postulated.     

Sympodial and monopodial branching inAcer (Aceraceae): Evolutionary trend and ecological implications

The evolutionary trend and its ecological implications in sympodial and monopodial branching patterns has been investigated in 20 JapaneseAcer spp. through comparison of shoot tip abortion and terminal bud formation. The genus is divided into two species groups according to its branching pattern, one (6 species) predominantly exhibiting sympodial branching with frequent monopodial branching in short shoots (sympodial species), and the other (14 species) exhibiting only monopodial branching (monopodial species). The early ontogeny of leaf and bud scales is described. Despite the difference in branching patterns, the bud scales of terminal buds are essentially the same in having a leaf base developed to function as a protecting organ. In all the sympodial species, during the abortion of a sympodium shoot tip, one or two pairs of primordia were found to occur on the apex, and later wither. These primordia resemble bud scales of terminal buds in their ontogeny and morphology, and appear to be rudimentary. It is suggested that a rudimentary terminal bud develops together with the establishment of sympodial branching, and that sympodial branching has originated from monopodial branching. Based on this proposed evolutionary trend, it is suggested thatAcer has moved from less shady habitats into shady habitats with monopodial branching (advantageous for vertical growth) changing into sympodial branching (advantageous for lateral spread).     

Typical cyperoid reproductive structures in Lipocarpha humboldtiana and Ascolepis brasiliensis (Cypereae – Cyperoideae – Cyperaceae): New evidence from a development perspective

Ascolepis and Lipocarpha, Cyperaceae, have highly reduced reproductive structures and hypogynous scales that are controversially appreciated. Because of this, flowers and spikelets and, thus, inflorescences have been interpreted in different ways, which, in turn, has led to placing the two genera in different tribes. Some authors interpret spikelets in Ascolepis and Lipocarpha as many-flowered and the so-called hypogynous scales as homologous to the lateral scales of Hypolytrum, or consider these scales comparable to a Scirpus bristle. However, many other authors consider spikelets in Lipocarpha and Ascolepis to be a result of a reductional process from a many-flowered cyperoid spikelet to a single-flowered spikelet, where the adaxial and abaxial hypogynous scales may be seen as the respective prophyll and glume of the reduced spikelets. The latest molecular phylogenies of Cyperaceae show both genera nested in the Cyperus clade, forming, in turn, a clade together with the rest of the Cypereae genera having single-flowered spikelets and hypogynous scales. Alternatively, based on this, the scales of uncertain homology that accompany the Ascolepis and Lipocarpha flower might be seen as special reproductive formations, representing a synapomorphy of such clade, instead of relicts of an ancestor with many-flowered spikelets. In view of this, freshly collected inflorescences of Lipocarpha humboldtiana Nees and Ascolepis brasiliensis (Kunth) Benth. ex C. B. Clarke were studied using light and scanning electron microscopy, with the aim of elucidating the nature of the controversially interpreted reproductive structures of these species from a development perspective. Results show that the “hypogynous scales” simply represent vestigial structures derived from the reduction of typical cyperoid spikelets, rather than a perianth part or specialized formations emerging as evolutionary novelties. Two scales are typically generated in both species, one being abaxial and the other adaxial, homologous to a glume and a prophyll, respectively, which contrasts with the eprophyllate condition so far attributed to Ascolepis. In both cases, the inflorescence is a spike of reduced spikelets, and the floral development in L. humboldtiana and in A. brasiliensis follows the general ontogenetic pattern observed in Cyperoideae. These characteristics support the inclusion of both genera in the Cypereae tribe.     

对小麦顶生小穗的初步研究

1.顶生小穗的护颖具有特殊的形态,第二护颖常为小花外稃状,腋内有时还保留着雌雄蕊或内稃残余。说明其不稳定和可变的本质。2.顶生小穗具特殊的坐落位置,其小穗轴与主穗轴一致。顶生小穗原始体发生在穗生长锥顶端,其下无苞原始体,长成后也无小穗领。其护颖和小花外稃与侧生小穗下的小穗领呈严格连续互生状态。说明其一次轴的渊源。3.顶生小穗护颖腋内可长出小穗,小花也可代之以小穗,护颖和小花外稃有时以苞片的形式保留于新侧生小穗外侧。新顶生小穗的护颖来自小花外稃。说明顶生小穗护颖腋内的退化花芽、外稃腋内的小花与侧生小穗都是花序一次轴上的二次轴分枝。4.顶生小穗产生小穗的变异严格按自下而上的顺序进行,与原侧生小穗有严格的连续性。5.事实证明,顶生小穗是一次轴花序,它属于穗状花序顶端的可变部分。     

Barley spikelet culture: An effective tool for the analysis of biochemical events in flower development

Immature detached barley spikelets were cultured in wheat spikelet medium. Fertility of cultured barley spikelets was similar to that of cultured wheat spikelets. Barley anther development within cultured spikelets was retarded relative to in planta, but viability of developing pollen, as determined by a fluorochromatic reaction, was similar in vitro and in planta. Protein synthesis by anthers in developing barley spikelets in vitro was maximal during stage 2, when pollen grains were bicellular, and declined as pollen matured. Barley spikelet culture is an effective tool for the analysis of biochemical events in flower development.Abbreviations FRC fluorochromatic reaction - WSM wheat spikelet medium     

苔草属植物的返祖两性小穗

本文报告了大别苔(Carex dabiecnsis)和白鳞苔草(C.polyschoena)的返祖小穗。这些返祖小穗中的类型之一具有与篙草属(Kobresia)以及Schoenoxiphium属完全相似的两性结构小穗,在这一类型的小穗中,它们生有一雌花及其在上的雄花。如图所示,大别苔和白鳞苔草中第(3)类型的返祖小穗和Schoenoxiphium属以及嵩草属的第1、2类型的小穗基本上没有区别,这些返祖小穗的发现证明果胞是苔草属的真正小穗,而且可以证明现存苔草的单性小穗是起源于具两性小穗的祖先。因此,可以相信具有两性结构的返祖小穗的发现进一步揭示了苔草亚科中Schoenoxiphium属,蒿草属,钩轴莎属和苔草属之间的系统发育的亲缘关系。     

The role of teosinte glume architecture (tga1) in coordinated regulation and evolution of grass glumes and inflorescence axes

? Hardened floral bracts and modifications to the inflorescence axis of grasses have been hypothesized to protect seeds from predation and/or aid seed dispersal, and have evolved multiple times independently within the family. Previous studies have demonstrated that mutations in the maize (Zea mays ssp. mays) gene teosinte glume architecture (tga1) underlie a reduction in hardened structures, yielding free fruits that are easy to harvest. It remains unclear whether the causative mutation(s) occurred in the cis-regulatory or protein-coding regions of tga1, and whether similar mutations in TGA1-like genes can explain variation in the dispersal unit in related grasses. ? To address these questions TGA1-like genes were cloned and sequenced from a number of grasses and analyzed phylogenetically in relation to morphology; protein expression was investigated by immunolocalization. ? TGA1-like proteins were expressed throughout the spikelet in the early development of all grasses, and throughout the flower of the grass relative Joinvillea. Later in development, expression patterns differed between Tripsacum dactyloides, maize and teosinte (Z. mays ssp. parviglumis). ? These results suggest an ancestral role for TGA1-like genes in early spikelet development, but do not support the hypothesis that TGA1-like genes have been repeatedly modified to affect glume and inflorescence axis diversification.     

Studies on the relationship between air temperature and the differentiation of young spikes of winter wheat in Beijing district

In these studies the optimum temperature indices for spikelet differentiation were found. The critical period determining the number of spikelets on a spike lies between the single ridge stage and the stage of glume differentiation. During this period a daily temperature below 7.5°C is favourable for differentiation of further spikelets. The processes of differentiation of wheat spikes need certain accumulated temperatures for a mean daily temperature above 0°C. The relationship between the rate of spikelet differentiation and temperature during the differentiation period, and the of these periods are discussed. According to the effect of climate in early spring on the number of differentiated spikelets of winter wheat, three climatic types in early spring are suggested.     

COMPARATIVE ANATOMY OF THE SPIKELET CALLUS OF ERIOCHLOA,BRACHIARIA, AND UROCHLOA (POACEAE: PANICEAE: SETARIINEAE)

Anatomical investigation of the spikelet base in Eriochloa reveals a nonvascularized cup-shaped structure composed of large parenchyma cells surrounding a column of tissue that is continuous with the pedicel. The cup and column are fused only at the base of the cup. No vestigial stelar node occurs in either the cup or the column. The stele branches above the cup at the rachillar node of the second glume. Therefore, the cup-shaped callus characteristic of Eriochloa appears to be formed entirely of parenchymatous tissue and is not derived from the first glume as previously interpreted. The bead of callus at the base of the spikelet in some species of Brachiaria also comprises an unbranched stele surrounded by parenchyma. The vascular tissue branches and enters the first glume at the apex of the bead. The spikelet base of Urochloa has a distinct first glume, nodal complex, and no callus parenchyma. Thus, the cup-shaped callus of Eriochloa and the callus bead of Brachiaria appear to be structurally similar.     

An architectural model for Eleocharis: morphology and development of Eleocharis cellulosa (Cyperaceae)

Species of Eleocharis are prominent in aquatic and wetland habitats and serve as models for study of physiological adaptations to aquatic environments. The genus has an unusual morphology because the major photosynthetic organ is the stem. In order to define an architectural model for the genus to understand the evolution of this morphology, we examined mature morphology and development of E. cellulosa in living and fixed material using light and scanning electron microscopy. Eleocharis cellulosa has sympodial, vertical shoots that produce the photosynthetic culms and horizontal shoots that mix monopodial and sympodial development. Each sympodial unit produces three bracts, an elongated photosynthetic internode, then a fourth bract and an inflorescence that either aborts on vegetative culms or expands on reproductive culms. On each sympodium, the first bract subtends a precocious axillary bud that reiterates the sympodial unit; the second bract subtends a bud that develops the horizontal shoot. In both horizontal and vertical shoots, the internode below the second bract is produced by both the second bract and the renewal shoot. Sympodial growth is present in seedlings. In other species of Eleocharis, the structure of the sympodial unit is conserved but morphological diversity develops from variation in horizontal shoot growth.     

Flower or spikelet? Understanding the morphology and development of reproductive structures in Exocarya (Cyperaceae, Mapanioideae, Chrysitricheae)

Fundamental questions of floral morphology remain unresolved in the grasslike monocots in order Poales, including what constitutes a flower and what constitutes a spikelet. The mapaniid sedges have particularly complex spikeletlike structures, variously interpreted as clusters of flowers or spikelets. Recent phylogenetic studies of Cyperaceae have identified the mapaniid clade as sister to the rest of the family, but the homology of mapaniid reproductive units (RUs) and spikeletlike units (SLUs) to other sedge flowers and spikelets is unclear. We examined reproductive development in the mapaniid Exocarya sclerioides. Inflorescence branches terminated in a SLU with bracts and 1-4 RUs. RUs had four small leaflike structures (LLSs): two lateral LLSs, each associated with a stamen, an abaxial LLS associated with a stamen, and an adaxial LLS. The gynoecium terminated the RU. All RUs were axillary to bracts, and unexpanded bracts and RUs were produced beyond expanded RUs, so SLUs were racemose. RUs developed from a single primordium that initiated two lateral LLSs, then two lateral stamens, then the gynoecium. Initiation of the abaxial LLS and stamen and the adaxial LLS followed. We hypothesize that the RU is a sympodial branch that terminates in a hermaphroditic flower with two stamens and a gynoecium; the two lateral LLSs are halves of a deeply divided prophyll.     

Studies on Morphological Evolution,Classification and Distribution of the Triticeae in China

_{The classification and the relationships among the genera of Chinese Triticeae were studied based on morphological characters with reference to geographical distribution and habitat conditions. The spike of Triticeae might have been derived from a panicled inflorescence like that in the Bromeae through a racemose inflorescence like the one in the Brachypodieae. There might be three evolutionary lines in the tribe. 1. Pedicels of the panicled inflorescence have become short and bracts decreased in size, which has resulted in a panicled spike with indefinite spikelets or false solitary spikelets at each node of rachis. The middle ribes of both glumes and lemmas and rachilla are not in a single plane. 2. A simple spike with usual solitary spikelets at each node of rachis has been derived from the raceme. The middle ribe of both glumes and lemmas and rachilla are in a single plane. 3. A cymose spike with 3-spikelets at each node of rachis has evolved from the cymose panicle. The glume on the central spikelet is behind the lemma, while those on the lateral spikelets are on lateral sides of the lemmas. From what we have described above Triticeae may be divided into three subtribes: Elyminae, Triticinae and Hordeinae. Then according to the morphological characters of glume, lemma and other organs as well as the habitats and distribution, the native and introduced triticeous plants are classified into 13 genera (Leymus, Elymus, Roegneria, Elytrigia, Aegilops, Triticum, Agropyron, Eremopyrum, Secale, Haynaldia, Psathyrostachys, Hordeum and Hystrix) and their relationships are also discussed meanwhile.}     

Floret position in Costularia (Cyperaceae): a new interpretation

The spikelet of Costularia has been interpreted as comprising proximal, sterile glumes followed by two larger fertile glumes that subtend respectively, a lower male and an upper bisexual floret. The terminal uppermost glume, with two keels and therefore resembling a prophyll, was empty. Studies of developmental stages of spikelets from Zimbabwe (Chimanimani Mts.) have revealed that the terminal glume envelops the bisexual floret and becomes empty only with maturation of the ovary.     

Understanding spikelet orientation in Paniceae (Poaceae)

Spikelet structure and grouping are key characters to identify grasses. Here we tested the possibility that spikelet pairs, a distinctive morphological structure of many Andropogoneae and Paniceae, are the starting point for a secondary single spikelet condition that can also explain the change of spikelet orientation among Paniceae genera. As a first approach, we studied the inflorescence development of Paspalum simplex, P. stellatum, and Axonopus sufultus to clarify the origin of the spikelet orientation and other basic homologies. The results support that solitary spikelets of A. suffultus are homologous to the subsessile spikelets of P. simplex and that solitary spikelets of P. stellatum are homologous to the pedicellate spikelet of P. simplex. This last homology supports that spikelet orientation results from a differential reduction/abortion of either the pedicellate or the subsessile spikelet primordia. We also discuss the possibility that the RAMOSA and polar auxin pathways could play a role in the abortion of the lateral subsessile spikelets in P. stellatum. However, the apical meristem inhibition observed in A. suffultus and P. stellatum seems to depend on a very different genetic control, suggesting that the single spikelet condition is homoplasic within Paniceae and derived from at least two different genetic mechanisms.     

Modeling Nymphoides architecture: A morphological analysis of Nymphoides aquatica (Menyanthaceae)

? Premise of the study: Species in the aquatic genus Nymphoides have inflorescences that appear to arise from the petioles of floating leaves. The inflorescence-floating leaf complex can produce vegetative propagules and/or additional inflorescences and leaves. We analyzed the morphology of N. aquatica to determine how this complex relates to whole plant architecture and whether whole plant growth is sympodial or monopodial. ? Methods: We used dissections, measurements, and microscopic observations of field-collected plants and plants cultivated for 2 years in outdoor tanks in south Florida, USA. ? Key results: Nymphoides aquatica had a submerged plagiotropic rhizome that produced floating leaves in an alternate/spiral phyllotaxy. Rhizomes were composed of successive sympodial units that varied in the number of leaves produced before the apex terminated. The basic sympodial unit had a prophyll that subtended a renewal-shoot bud, a short-petioled leaf (SPL) with floating lamina, and an inflorescence; the SPL axillary bud expanded as a vegetative propagule. Plants produced either successive basic sympodial units or expanded sympodia that intercalated long-petioled leaves between the prophyll and the SPL. ? Conclusions: Nymphoides aquatica grows sympodially, forming a rhizome composed of successive basic sympodia and expanded sympodial units. Variations on these types of sympodial growth help explain the branching patterns and leaf morphologies described for other Nymphoides species. Monitoring how these two sympodial phases are affected by water depth provides an ecologically meaningful way to assess N. aquatica's responses to altered hydrology.