Development of the mixed inflorescence in Zea diploperennis litis, Doebley & Guzman (Poaceae)

Development of the mixed inflorescence in Zea diploperennis Iltis, Doebley & Guzman (Poaceae) Mixed inflorescences of diploperennial teosinte, which terminate the main branches of the plant, arise in the same fashion as tassel spikes. The apical meristem produces bracts in a decussate arrangement. A single axillary bud primordium is initiated in the axil of each bract. Growth of the bract is retarded as the bud enlarges and divides longitudinally into two separate spikelet primordia. The paired spikelets running in two ranks on either side of the inflorescence primordium produce the four-rowed condition typical of teosinte tasselS. In the transition region between male and female portions of the inflorescence, development of the pedicellate spikelet of each spikelet pair is arrested at an early ontogenetic stage. Continued growth of the sessile spikelet and associated rachis flaps destroy the remnants of the arrested spikelet in basal portions of the inflorescence. A similar abortion of the lower floret of the sessile spikelet results in a single pistillate floret per node at anthesis. These results provide further support for the hypothesis that a tassel-like mixed inflorescence of teosinte is ancestral to the maize ear.     

Analysis of inflorescence organogenesis in eastern gamagrass, Tripsacum dactyloides (Poaceae): the wild type and the gynomonoecious gsf1 mutant

Inflorescence organogenesis of a wild-type and a gynomonoecious (pistillate) mutant in Tripsacum dactyloides was studied using scanning electron microscopy. SEM (scanning electron microscope) analysis indicated that wild-type T. dactyloides (Eastern gamagrass) expressed a pattern of inflorescence organogenesis that is observed in other members of the subtribe Tripsacinae (Zea: maize and teosinte), family Poaceae. Branch primordia are initiated acropetally along the rachis of wild-type inflorescences in a distichous arrangement. Branch primordia at the base of some inflorescences develop into long branches, which themselves produce an acropetal series of distichous spikelet pair primordia. All other branch primordia function as spikelet pair primordia and bifurcate into pedicellate and sessile spikelet primordia. In all wild-type inflorescences development of the pedicellate spikelets is arrested in the proximal portion of the rachis, and these spikelets abort, leaving two rows of solitary sessile spikelets. Organogenesis of spikelets and florets in wild-type inflorescences is similar to that previously described in maize and the teosintes. Our analysis of gsf1 mutant inflorescences reveals a pattern of development similar to that of the wild type, but differs from the wild type in retaining (1) the pistillate condition in paired spikelets along the distal portion of the rachis and (2) the lower floret in sessile spikelets in the proximal region of the rachis. The gsf1 mutation blocks gynoecial tissue abortion in both the paired-spikelet and the unpaired-spikelet zone. This study supports the hypothesis that both femaleness and maleness in Zea and Tripsacum inflorescences are derived from a common developmental pathway. The pattern of inflorescence development is not inconsistent with the view that the maize ear was derived from a Tripsacum genomic background.     

毛竹的花序发育研究

通过野外观察和石蜡切片技术研究了毛竹(Phyllostachys edulis)的花序发育进程。研究结果表明:毛竹的花序为续次发生的假花序,以小穗为单元,4~13个不等,偏向一侧排列(似扫帚状)的小穗组成长约8.01 cm的复穗状花序;当花序伸长至4~5 cm时,形成侧芽结构,小穗原基开始发育,形成各级小穗,直至顶生小穗、侧生小穗出现;当花序伸长至8~10 cm时,颖花原基形成并开始发育,最终形成3个雄蕊和1个雌蕊构成的小花。花序形成初期(5月中旬至6月),苞片紧裹主轴,顶端具缩小叶;随着分蘖小穗的生长和小花开放,植株叶片变黄,整个花序变为褐色,进入种实发育成熟阶段。本文首次报道了毛竹花序的发育进程,进一步丰富了竹类生殖生物学的研究内容,为竹亚科及禾本科的生殖生物学研究积累了丰富的材料。     

A Taxonomical Study on the Genus Shibataea Makino

Shibataea Makino is a genus of Subfam. Bambusoideae, with 8 species, distributed in Southeast China and Southwest Japan. In China wild plants of the genus are found in Fujian, Jiangxi, Zhejiang, Jiangsu and Anhui provinces, especially in Fujian and Zhejiang. The genus is also cultivated in parks of Guangzhou, Teibei and some other gardens. Raches of inflorescences in genera Semiarundinaria, Brachystachyum, Phyllostachys and Shibataea have many branches, even secondary branches. A large bract is often present at the base of each branch, and a prophyll in the axil of the bract in Tribe Shibataeeae Nakai. Moreover, an inflorescence is composed of numerous dense spikelets. This type of inflorescence may be considered primitive. The genera Indosasa and Sinobambusa are of more stamens (6 in the former and 3 or 4, 5 in the latter) than in the genera Semiarundinaria and Brachystachyum (only 3), and their inflorescences are very simple with fewer spikelets and raches,without the large bract. This type of inflorescence may be considered more advanced.     

Inflorescence development in a new teosinte: Zea nicaraguensis (Poaceae)

Inflorescence development in a newly discovered teosinte, Zea nicaraguensis (Poaceae), from Nicaragua has been investigated using scanning electron microscopy (SEM). The SEM examination revealed that the pattern of both male and female inflorescence development was similar to previously described inflorescence in other Zea taxa. Branch primordia were initiated acropetally in a distichous pattern along the rachis of male and female inflorescences. Spikelet pair primordia bifurcated into pedicellate and sessile spikelet primordia. Predictably, pedicellate spikelet development was arrested and aborted in the female teosinte inflorescence. Organogenesis of functional spikelets and florets was similar to that previously described in maize and teosintes. The results were consistent with our hypothesis that both femininity and masculinity share a common mechanism of inflorescence development in Zea and Tripsacum and are in accord with a putative common mechanism of sex determination in the Andropogoneae. Interestingly, this population of teosinte, unique in its ability to grow in water-logged soils, showed a stable pattern of early inflorescence development. Our results also revealed the uncharacteristic presence of inflorescence polystichy in this population of Zea nicaraguensis. We propose this novel phenotypic variation raises the possibility that a domestic evolution of polystichy in maize was enabled by an occasional polystichous phenotypic in teosinte.     

Unisexual flowers as a robust synapomorphy in Cariceae (Cyperaceae)? Evidence for bisexual flowers in Schoenoxiphium

Cariceae, the largest tribe within Cyperaceae, comprises about 2000 species in five genera. Cariceae is usually considered to be distinct from other Cyperaceae by the presence of exclusively unisexual flowers and by the arrangement of the pistillate flowers in single-flowered spikelets that are enclosed by the flask-like spikelet prophyll (utricle or perigynium). The nature of several morphological features of the Cariceae inflorescence remains controversial. The staminate reproductive units, as well as earlier reported bisexual reproductive units in Schoenoxiphium have been considered to be reduced partial inflorescences, or flowers. Aims of this study are to test both interpretations, based on a floral ontogenetic investigation. Moreover, for the first time, detailed SEM micrographs are presented of the inflorescence and floral development and of bisexual flowers in Schoenoxiphium. We propose that ‘inhibition of bisexuality’ is a more robust synapomorphy in Cariceae than ‘presence of only unisexual flowers’.     

Variability of the inflorescence morphology of Carex spicata (Cyperaceae) and its implication to taxonomy

Plants growing in different kinds of habitats are expected to show high morphological plasticity. Carex spicata Huds. occurs in different plant communities and shows distinct morphological variability of the inflorescences. Field observations carried out in different plant communities permitted us to hypothesize that the morphological variability of C. spicata inflorescences may to some degree be the result of the habitat. The objective of this study was to test that hypothesis for C. spicata by collecting inflorescences and measuring their morphological features from several populations in each of six plant communities: Agropyro‐Urticetum dioiceae, Arrhenatheretum elatioris, Lolio‐Cynosuretum, Lolio‐Plantaginetum, Trifolio‐Agrimonietum, and Stellario‐Deschampsietum. The following inflorescence features were analyzed: length of the lowest spikelet, distance between two lowest spikelets, length of inflorescence, number of spikelets, and length proportion of the lowest spikelet to the distance between the two lowest spikelets. We found that all analyzed morphological characters differed significantly among the plant communities. Furthermore, we found significant differences among populations within the six plant communities. Moreover, C. spicata inflorescences often have morphological features (i.e. a less crowded inflorescence with a relatively large distance between the two lowest spikelets) similar to the closely related species C. muricata L. Thus, the limited diagnostic value of inflorescence crowding, especially in vegetative or early fruiting phases, suggests that the taxonomic importance of this character should be reconsidered.     

DEVELOPMENT OF TASSEL SEED 2 INFLORESCENCES IN MAIZE

The normal pattern of maize floral development of staminate florets on the terminal inflorescence (tassel) and pistillate florets on the lateral inflorescences (ears) is disrupted by the recessive mutation tassel seed 2. Tassel seed 2 mutant plants develop pistillate florets instead of staminate florets in the tassel. In addition, the ears of tassel seed 2 plants display irregular rowing of kernels due to the development of the normally suppressed lower floret of each spikelet. The morphology of tassel and ear florets of the recessive maize mutant tassel seed 2 has been compared to those of wild-type maize through development. We have identified the earliest stages at which morphological signs of sex differentiation are evident. We find that sex determination occurs during the same stage on tassel and ear development. Early postsex determination morphology of florets in wild-type ears and in tassel seed 2 tassels and ears is identical.     

Organogenesis of Fascicled ear mutant inflorescences in maize (Poaceae)

The ontogeny of staminate tassels and pistillate ears in the maize mutant Fascicled ear was examined using scanning electron microscopy. The normal pattern of inflorescence development is perturbed by the Fascicled ear mutation at the transition stage. The Fascicled ear mutation promotes the development of an abnormal transition stage axis that is both shorter and broader than the wild type. The inflorescence apical meristem then undergoes a bifurcation, and two inflorescence axes arise in place of a single axis. Each derived inflorescence apical meristem may undergo a similar perturbation sequence. This expression of the Fascicled ear mutation may be repeated one to several times, which leads to the development of a fascicled pistillate inflorescence and a fascicled central spike in the staminate inflorescence. The apical meristems of some tassel branches are also bifurcated. Subsequent organogenesis during paired-spikelet and floral development in Fascicled ear plants follows the pattern of normal maize. However, triplet spikelets are occasionally observed. The organogenic disruption by the Fascicled ear mutation that we describe will aid genetic and molecular analysis on the regulation of inflorescence development in maize and other members of the genus Zea.     

Vegetative Proliferation and Secondary Proliferated Inflorescences Development in Grass Ischaemum barbatum Retz

We report the vegetative proliferation and new phenomenon of ＂secondary proliferated inflorescences＂ In the grass Ischaemum barbatum Retz, as determined by anatomical analysis of prepared sections of inflorescences. Leaves and shoots could be developed from the original splkelets of Inflorescences and plantlets developed when these shoots were transplanted to moist soil. ＂Secondary proliferated Inflorescences＂ is the first name here because some inflorescences that developed inadequacy are grown from the splkelet on the mother Inflorescence. Our investigation showed that this form of vegetative proliferation and secondary proliferated inflorescences development of L barbatum has arisen following late autumn fires of the previous year. It Is suggested that the sudden onset of a fire could lead to a hormone Imbalance or a chemical induction, which results in ephemeral vegetative proliferation even secondary proliferated inflorescences development in wild populations.     

Phyllotaxis in the Oil Palm: Arrangement of Male/Female Florets along the Spikelets

The paper continues an earlier study of the geometry of inflorescencestructures in the oil palm in which geometry is measured interms of Equivalent Phyllotaxis Index (E.P.I.). In this casethe phyllotaxis of male and female florets along their respectivespikelets is considered. Regardless of the spikelet positionon the inflorescence or the palm age the very small male floretshave a higher E.P.I. than the large female structures and acompletely different apparent parastichy arrangement. TheseE.P.I. estimates seem to be independent of the age of palmsfrom which inflorescences and hence spikelets are sampled. However,there is considerable variation in phyllotaxis within bunches,E.P.I. being lower on spikelets sampled toward the base of theinflorescence and increasing in a more or less linear mannerin spikelets sampled at the tip; this pattern is not so definiteon male spikelets. The results are discussed in relation toother more simple measurements of spikelet architecture.     

Diversity,systematics, and evolution of Cynodonteae inflorescences (Chloridoideae – Poaceae)

The species of the Cynodonteae tribe show great morphological diversity in their reproductive structures. Previous studies where inflorescences were comparatively analysed in the context of phylogeny have shown that although grass inflorescences seem to be excessively variable, there are certain aspects of inflorescences that store relevant information on the evolution and systematics in Poaceae. We have analysed and compared the inflorescence structures of species belonging to the Hilariinae, Monanthochloinae, Scleropogoninae, and Muhlenbergiinae subtribes. Considering the most relevant morphological characters, the most recurrent types of inflorescences in the lineage were determined by means of a principal coordinates analysis. To understand the evolution of inflorescence morphology, ancestral reconstructions of inflorescence characters were performed using the Bayesian inference method. The results obtained demonstrate that the processes of homogenization and truncation might account for the diversity observed in adult inflorescences. Five different types of inflorescences were identified out of 36 theoretical possibilities. Amongst these, inflorescence type 1 (panicle of spikelets, with a terminal spikelet, non-homogenized, and bearing third- or higher-order branches) was found to be the most frequent in the studied group. Ancestral reconstructions of morphological characters allowed us to suggest that the ancestor of the group might have had an inflorescence with the form of a raceme of spikelets, non-truncated and bearing first-order branches. More complex inflorescences bearing no terminal spikelets and having branches of higher order might have diverged this lineage.     

Morphogenesis of Pistillate Flowers of Cercidiphyllum japonicum(Cercidiphyllaceae)

Floral morphogenesis and the development of Cercidiphyllum japonicum Sieb.et Zucc.were observed by scanning electronmicroscopy(SEM).The results showed that the pistillate inflorescences were congested spikes with the flowers arrangedopposite.Great differences between the so-called"bract"and the vegetative leaf were observed both in morphogenesis andmorphology.In morphogenesis,the"bract"primordium is crescent-shaped,truncated at the apex and not conduplicate,has no stipule primordium at the base but does have some inconspicuous teeth in the margin that are not glandular.Theleaf primordium is triangular,cycloidal at the apex,conduplicate,has two stipule primordia at the base,has one gland-toothat the apex occurring at first and some gland-teeth in the margin that occur later.In morphology,the"bract"is also differentto the vegetative leaf in some characteristics that were also illustrated in the present paper.Based on the hypothesis thatthe bract is more similar to the vegetative leaf than the tepal,we considered that the so-called"bract"of C.japonicum mightbe the tepal of the pistillate flower in morphological nature.Therefore,each pistillate flower contains a tepal and a carpel.We did not find any trace of other floral organs in the morphogenesis of the pistillate flower.Therefore we consideredthat the unicarpellate status of extant Cercidiphyllum might be to highly reduce and advance characteristics that make theextant Cercidiphyllum isolated from both fossil Cercidiphyllum-like plants and its extant affinities.     

Morphogenesis of Pistillate Flowers of Cercidiphyllum japonicum （Cercidiphyllaceae）

Floral morphogenesis and the development of Cercidiphyllumjaponicum Sieb. et Zucc. were observed by scanning electron microscopy （SEM）. The results showed that the pistillate inflorescences were congested spikes with the flowers arranged opposite. Great differences between the so-called ＂bract＂ and the vegetative leaf were observed both in morphogenesis and morphology. In morphogenesis, the ＂bract＂ primordium is crescent-shaped, truncated at the apex and not conduplicate, has no stipule primordium at the base but does have some inconspicuous teeth in the margin that are not glandular. The leaf primordium is triangular, cycloidal at the apex, conduplicate, has two stipule primordia at the base, has one gland-tooth at the apex occurring at first and some gland-teeth in the margin that occur later. In morphology, the ＂bract＂ is also different to the vegetative leaf in some characteristics that were also illustrated in the present paper. Based on the hypothesis that the bract is more similar to the vegetative leaf than the tepal, we considered that the so-called ＂bract＂ of C.japonicum might be the tepal of the pistillate flower in morphological nature. Therefore, each pistillate flower contains a tepal and a carpel. We did not find any trace of other floral organs in the morphogenesis of the pistillate flower. Therefore we considered that the unicarpellate status of extant Cercidiphyllum might be to highly reduce and advance characteristics that make the extant Cercidiphyllum isolated from both fossil Cercidiphyllum-like plants and its extant affinities.     

Observations of inflorescence patterns and flower sex in a population of Sagittaria papillosa Buch. (Alismataceae)

Sagittaria papillosa Buch. is monoecious with unisexual flowers, pistillate below, staminate above, typically with an unbranched scape. A large population with unusual numbers of staminate and bisexual flowers on the lowest whorl of the inflorescence and many particles was quantitatively evaluated. First-formed inflorescences had more staminate and bisexual flowers than those produced later. Branched scapes were predominantly found to be the second inflorescence produced by a given plant. Genetic crosses between flowers on recemes and panicles produced no branched inflorescences. When grown under greenhouse conditions all tested plants had racemes with pistillate flowers in the lower whorls and staminate ones above. Data from soil parameters, daylengths and air temperatures are compared to reported information on modification of flower sexuality by these factors.     

An experimental study of the interaction between the dwarf palm (Chamaerops humilis) and its floral visitor Derelomus chamaeropsis throughout the life cycle of the weevil

Palm pollination can be quite diverse but has been poorly studied. This paper describes the life cycle of Derelomus chamaeropsis, a Coleoptera that inhabits the inflorescences of the Mediterranean dwarf palm Chamaerops humilis. D. chamaeropsis is specific to Chamaerops inflorescences, where it eats pollen and the rachis of inflorescences on pistillate plants. They usually lay eggs only on staminate inflorescences where larvae develop and bore into the inflorescence rachis. Larvae do not develop on pistillate inflorescences, except for cases with almost no fruit development. Pistillate plants can thus protect themselves from weevil predation. When visiting pistillate inflorescences, weevils can feed on rachis but usually do not find the brood place reward. Pollination is thus by deceit and weevils should be selected to avoid pistillate inflorescences. D. chamaeropsis pupate within the rachis of staminate inflorescences, but disperse before collecting pollen, thus staminate plants do not have an individual advantage in breeding weevils. However, because larvae develop on dead tissues, the costs of larval development are likely to be low for the plant. This study provides a new example of pollination symbiosis where the pollinator develops on the plant it pollinates, and illustrates how the evolutionary functioning of such relationships can be diverse.     

barren inflorescence2 regulates axillary meristem development in the maize inflorescence

Organogenesis in plants is controlled by meristems. Shoot apical meristems form at the apex of the plant and produce leaf primordia on their flanks. Axillary meristems, which form in the axils of leaf primordia, give rise to branches and flowers and therefore play a critical role in plant architecture and reproduction. To understand how axillary meristems are initiated and maintained, we characterized the barren inflorescence2 mutant, which affects axillary meristems in the maize inflorescence. Scanning electron microscopy, histology and RNA in situ hybridization using knotted1 as a marker for meristematic tissue show that barren inflorescence2 mutants make fewer branches owing to a defect in branch meristem initiation. The construction of the double mutant between barren inflorescence2 and tasselsheath reveals that the function of barren inflorescence2 is specific to the formation of branch meristems rather than bract leaf primordia. Normal maize inflorescences sequentially produce three types of axillary meristem: branch meristem, spikelet meristem and floral meristem. Introgression of the barren inflorescence2 mutant into genetic backgrounds in which the phenotype was weaker illustrates additional roles of barren inflorescence2 in these axillary meristems. Branch, spikelet and floral meristems that form in these lines are defective, resulting in the production of fewer floral structures. Because the defects involve the number of organs produced at each stage of development, we conclude that barren inflorescence2 is required for maintenance of all types of axillary meristem in the inflorescence. This defect allows us to infer the sequence of events that takes place during maize inflorescence development. Furthermore, the defect in branch meristem formation provides insight into the role of knotted1 and barren inflorescence2 in axillary meristem initiation.     

Class II tassel seed mutations provide evidence for multiple types of inflorescence meristems in maize (Poaceae)

The tassel seed mutations ts4 and Ts6 of maize cause irregular branching in its inflorescences, tassels, and ears, in addition to feminization of the tassel due to the failure to abort pistils. A comparison of the development of mutant and wild-type tassels and ears using scanning electron microscopy reveals that at least four reproductive meristem types can be identified in maize: the inflorescence meristem, the spikelet pair meristem, the spikelet meristem, and the floret meristem. ts4 and Ts6 mutations affect the fate of specific reproductive meristems in both tassels and ears. ts4 mutants fail to form spikelet meristems from spikelet pair meristems. Ts6 mutants are delayed in the conversion of certain spikelet meristems into floret meristems. Once floret meristems are established in both of these mutants, they form florets that appear normal but fail to undergo pistil abortion in the tassel. The abnormal branching associated with each mutant is suppressed at the base of ears, permitting the formation of normal, fertile spikelets. The classification of the different types of reproductive meristems will be useful in interpretation of gene expression patterns in maize. It also provides a framework for understanding meristem functions that can be varied to diversify inflorescence architectures in the Gramineae.