Types of enrichment axes in Poaceae

The objective of this review work is to characterize the enrichment axes in Poaceae, especially integrating into that analysis of those species with basal or subterranean cleistogamous spikelets. We recognize five types of enrichment axes: paraclades of the unit of inflorescence (UIF), paraclades of the trophotagma (TT) with exposed UIF; paraclades of the trophotagma with not exposed UIF; subterranean paraclades on short rhizomes and subterranean paraclades upon plagiotropic axes of long internodes. According to the enrichment axes, we differentiate six types of synflorescences. The different types of enrichment axes and synflorescences types are characterized; their differences determine in some cases the existence of fruit heteromorphism.     

濒危植物庙台槭生殖生物学研究1.花序和花的形态及发育

庙台械的花序为有限花序,由一顶花和6—9枝侧花枝组成,属圆锥状聚伞花序。一个花序共有14—29朵花,包括两性花、雄花和无性花三类花。根据花在花序上着生的位置,可分为三级。7月初,花序原基形成,在花序轴伸长的同时,侧面形成侧花枝轴原基。花序的顶花最早进行个体发育,随后是侧花枝顶花;侧花枝上同一级花的发育顺序则是从花序的下面向上进行。花器官发生时,花萼原基最先形成,然后是花瓣、雄蕊、心皮和胚珠。     

Inflorescence morphology, heterochrony, and phytogeny in the mimosoid tribes ingeae and acacieae (Leguminosae: Mimosoideae)

In earlier work (Grimes, 1992) on inflorescence morphology in the mimosoid tribes Ingeae and Acacieae I proposed that differences in inflorescence morphology result from three properties: the organization of components of the inflorescence and their relative positions; the hierarchical arrangement of the axes of the inflorescence and the position they assume in total tree architecture; and the heterochronic development of components of the inflorescence. Further work shows that the first two properties are better stated in terms of heterochrony; namely, that the organization of components of the inflorescence differs due to differences in timing of the development of organ systems and that the hierarchy of axes likewise differs due to heterochronic changes. Neither de novo origin of organs or organ systems nor suppression or loss of organs or organ systems accounts for the diversity in form. Observed heterochronic differences in the inflorescence structure may be divided into three types: spatial differences in the relationship between the unit inflorescence and the subtending leaf (hysteranthy); differences in the time of formation and/or the duration of whole axes; and changes in development pathways, leading to shoot dimorphism. These heterochronies are used as characters in a cladistic analysis, and it is shown that although some are homoplasious, many provide synapomorphies of clades of exemplars representing genera in the Ingeae and Acacieae.     

Requirement of the Auxin Polar Transport System in Early Stages of Arabidopsis Floral Bud Formation

The pin-formed mutant pin 1-1, one of the Arabidopsis flower mutants, has several structural abnormalities in inflorescence axes, flowers, and leaves. In some cases, pin1-1 forms a flower with abnormal structure (wide petals, no stamens, pistil-like structure with no ovules in the ovary) at the top of inflorescence axes. In other cases, no floral buds are formed on the axes. An independently isolated allelic mutant (pin1-2) shows similar phenotypes. These mutant phenotypes are exactly the same in wild-type plants cultured in the presence of chemical compounds known as auxin polar transport inhibitors: 9-hydroxyfluorene-9-carboxylic acid or N-(1-naphthyl)phthalamic acid. We tested the polar transport activity of indole-3-acetic acid and the endogenous amount of free indole-3-acetic acid in the tissue of inflorescence axes of the pin1 mutants and wild type. The polar transport activity in the pin 1-1 mutant and in the pin1-2 mutant was decreased to 14% and 7% of wild type, respectively. These observations strongly suggest that the normal level of polar transport activity in the inflorescence axes is required in early developmental stages of floral bud formation in Arabidopsis and that the primary function of the pin1 gene is auxin polar transport in the inflorescence axis.     

孟仁草的花序发育研究

在光学显微镜和扫描电镜下观察了禾本科(Poaceae)虎尾草属(Chloris Sw.)孟仁草(Chloris barbata Sw.)的花序发育过程,以寻找适于虎尾草群(Chloris group)分支分析的发育性状.结果发现了未见于成熟花序的23个发育性状.阐明盂仁草花序的本质是二级长侧枝包围平截的主轴构成指形花序.该类型花序仅见于单子叶植物和少数高度特化的双子叶植物.涉及花序分枝的分子遗传机制研究亟待开展.     

Structure of the Cyperaceae Inflorescence

This work presents the basics for interpreting the adult inflorescence structure in Cyperaceae. It provides an analysis of the variations of the synflorescence and inflorescence structure in the family. Three types of synflorescence may be recognized in this family: a synflorescence with a foliate stem, a terminal inflorescence and a variable number of lateral inflorescences; a synflorescence with a foliate stem and only the terminal inflorescence; and a synflorescence with a scape and a terminal inflorescence. Variations in the structure and form of the inflorescences are related to variations in inflorescence branching, inflorescence homogenization degree, presence or absence of the distal part of the inflorescence, phyllotaxis, inflorescence position, types of bracts and leaves subtending branches, elongation of inflorescence internodes and spikelet structure. These variations are correlated with some of the developmental processes that give origin to the inflorescence.     

Enhanced inflorescence development in bougainvillea "san diego red" by removal of young leaves and cytokinin treatments

Removal of young leaves and application of the cytokinin, N-benzyla-α-(tetrahydro-2H-pyran-2yl)-adenine promote inflorescence development in Bougainvillea “San Diego Red”. Defoliation greatly increased the amount of assimilate accumulated at the shoot tip 1 to 2 days after treatment. Cytokinin applications further increased the amount accumulated and this increase was apparent 4 days before morphological changes could be detected at the inflorescence axes. Short days promoted inflorescence development and also increased assimilate accumulation at the reproductive axes; thus, it is suggested that the role of short day induction in bougainvillea may be that of redirecting the flow of assimilates, perhaps by its influence on cytokinin synthesis and distribution.     

Water Deficit and Inflorescence Development in Zea mays L.--The Role of the Developing Tassel

In the normal pattern of development of Zea mays (cv. Iochief)a single mature female inflorescence is produced at node 7.A brief episode of water deficit at the time of terminal maleinflorescence initiation induced the subsequent developmentof two to three mature female inflorescences at nodes 5–7.This growth of the inflorescences at lower nodes was accompaniedby a marked inhibition of the growth of the terminal male inflorescence.Removal of either the developing terminal inflorescence or ofthe axillary inflorescence at node 7 at this time also promotedthe growth of the lower axillary inflorescences. The growthof these inflorescences was further stimulated by a period ofwater deficit when only the inflorescence at node 7 was removed,but removal of the male inflorescence abolished the capacityof these inflorescences to respond to the water deficit Excisionof the male inflorescence immediately before or immediatelyafter the period of water deficit produced the same response.It is concluded that this response of the lower axillary inflorescencesto water deficit is mediated through an effect on the developingterminal male inflorescence. Zea mays, water deficit, inflorescence development, tassel, correlative inhibition     

The inflorescence structure of Kummerowia (Leguminosae)

Inflorescences of Kummerowia are compound and the component axes appear to terminate in a flower. In order to clarify whether or not the flower is truly terminal, inflorescences of Kummerowia were studied organographically, ontogenetically and anatomically. Four inflorescence phyllomes are usually produced immediately below the seemingly terminal flower and appear to be borne on the same axis. The second phyllome subsequent to the lowest one is located at right angles to the lowest one, and the third and fourth ones located opposite each other and at right angles to the second. The lowest phyllome is sometimes undeveloped in K.stipulacea. Ontogenetic observation revealed the presence of two abortive apiceS. Anatomical observation revealed that these two abortive apices remain rudimentary in the flowering stage. On the basis of the arrangement of these phyllomes and the presence of the remnants of apices, the structure of the component inflorescence axis in Kummerowia is interpreted as follows: the component axis branches off a lateral axis, which is reduced entirely in length, from the axil of the lowest phyllome, and terminates in an abortive apex; the lateral axis in turn branches off one lateral axis of the next order, which is also reduced in length, from the axil of the second phyllome and terminates in an abortive apex; the lateral axis of the next order produces the third and fourth phyllomes and is terminated by a flower. The flower, which seems to terminate the component axis, is therefore axillary in origin. The axillary branch of the lowest phyllomes occasionally bears two lateral flowers. The branching system of the inflorescence of Kummerowia is identical with that of an inflorescence of Lespedeza cuneata. Kummerowia and Lespedeza are continuous in characteristics of the inflorescence, indicating the relationship between the inflorescence of Kummerowia and the pseudoraceme of Lespedeza.     

The distinctive features of ontogenetic processes of <Emphasis Type="Italic">Andromeda polifolia</Emphasis> L. (Ericaceae Juss.) in the European subarctic

The development cycle of the elementary unit of a shoot system (EUS) and of perennial axes consisting of these shoots in adult plants of Andromeda polifolia L. was investigated in the Kola Peninsula. It was established that EUS is a monocarpic shoot consisting of 14–26 metameres with three-year life cycle. The cycle consists of four phases: the initial intrabud (one year), vegetation (two to three months), intermediate intrabud with rudimentary inflorescence (one year), flowering and fruit-bearing (two to three months). Perennial skeletal axes (sympodia) consist of three to eight EUS. Development of the inflorescence and flowers is described. It is supposed that the large morphogenetic reserve of plants as numerous well developed capacious buds and their rapid development promote this species to northern regions where the vegetation period is short.     

宽叶金粟兰滇东北居群的繁殖生物学

对宽叶金粟兰（ＣｈｌｏｒａｎｔｈｕｓｈｅｎｒｙｉＨｅｍｓｌ）滇东北的野外居群和栽培植株花序发育、传粉、结实率、无性繁殖等观察试验的结果显示,宽叶金粟兰以种子繁殖为主,但有时可通过地下合轴茎顶端分支进行无性繁殖。其顶生、侧生花序均能正常开花结实,种子可萌发,在株丛周缘形成新个体。但其顶生花序与侧生花序的雄蕊异形,顶生花序花芽在地下分化形成,雄蕊具明显伸长的３个药隔和４个药室,不产香味,平均结实率６３％;侧生２、３级花序在直立枝条顶端叶腋中分化形成,无伸长药隔并仅具２个药室,在顶生花序果脱落后才开花结实,平均结实率７６％和８３％,略高于顶生花序。传粉实验观察证明其不需要昆虫传粉,为自花授粉可育,不同于本属其它植物。因此其雄蕊异型现象可能是传媒方式改变的结构简化适应     

Racemose inflorescences of monocots: structural and morphogenetic interaction at the flower/inflorescence level

Background

Understanding and modelling early events of floral meristem patterning and floral development requires consideration of positional information regarding the organs surrounding the floral meristem, such as the flower-subtending bracts (FSBs) and floral prophylls (bracteoles). In common with models of regulation of floral patterning, the simplest models of phyllotaxy consider only unbranched uniaxial systems. Racemose inflorescences and thyrses offer a useful model system for investigating morphogenetic interactions between organs belonging to different axes.

Scope

This review considers (1) racemose inflorescences of early-divergent and lilioid monocots and their possible relationship with other inflorescence types, (2) hypotheses on the morphogenetic significance of phyllomes surrounding developing flowers, (3) patterns of FSB reduction and (4) vascular patterns in the primary inflorescence axis and lateral pedicels.

Conclusions

Racemose (partial) inflorescences represent the plesiomorphic condition in monocots. The presence or absence of a terminal flower or flower-like structure is labile among early-divergent monocots. In some Alismatales, a few-flowered racemose inflorescence can be entirely transformed into a terminal ‘flower’. The presence or absence and position of additional phyllomes on the lateral pedicels represent important taxonomic markers and key features in regulation of flower patterning. Racemose inflorescences with a single floral prophyll are closely related to thyrses. Floral patterning is either unidirectional or simultaneous in species that lack a floral prophyll or possess a single adaxial floral prophyll and usually spiral in the outer perianth whorl in species with a transversely oriented floral prophyll. Inhibitory fields of surrounding phyllomes are relevant but insufficient to explain these patterns; other important factors are meristem space economy and/or the inhibitory activity of the primary inflorescence axis. Two patterns of FSB reduction exist in basal monocots: (1) complete FSB suppression (cryptic flower-subtending bract) and (2) formation of a ‘hybrid’ organ by overlap of the developmental programmes of the FSB and the first abaxial organ formed on the floral pedicel. FSB reduction affects patterns of interaction between the conductive systems of the flower and the primary inflorescence axis.     

A Mutation in the Arabidopsis TFL1 Gene Affects Inflorescence Meristem Development

We present the initial phenotypic characterization of an Arabidopsis mutation, terminal flower 1-1 (tfl1-1), that identifies a new genetic locus, TFL1. The tfl1-1 mutation causes early flowering and limits the development of the normally indeterminate inflorescence by promoting the formation of a terminal floral meristem. Inflorescence development in mutant plants often terminates with a compound floral structure consisting of the terminal flower and one or two subtending lateral flowers. The distal-most flowers frequently contain chimeric floral organs. Light microscopic examination shows no structural aberrations in the vegetative meristem or in the inflorescence meristem before the formation of floral buttresses. The wild-type appearance of lateral flowers and observations of double mutant combinations of tfl1-1 with the floral morphogenesis mutations apetala 1-1 (ap1-1), ap2-1, and agamous (ag) suggest that the tfl1-1 mutation does not affect normal floral meristems. Secondary flower formation usually associated with the ap1-1 mutation is suppressed in the terminal flower, but not in the lateral flowers, of tfl1-1 ap1-1 double mutants. Our results suggest that tfl1-1 perturbs the establishment and maintenance of the inflorescence meristem. The mutation lies on the top arm of chromosome 5 approximately 2.8 centimorgans from the restriction fragment length polymorphism marker 217.     

Inflorescence and floral development in Streptocarpus and Saintpaulia (Gesneriaceae) with particular reference to the impact of bracteole suppression

Floral development and inflorescence structure within Streptocarpus and Saintpaulia were investigated using Scanning Electron Microscopy (SEM). We discuss the structure and development of the pair-flowered cyme and the floral ontogeny found in the Gesneriaceae in a phylogenetic context with particular reference to an East African clade of Streptocarpus and Saintpaulia. Current phylogenetic hypotheses divide the caulescent East African Streptocarpus species into two distinct clades, in relation to which the position of Saintpaulia is not yet clear. Variation in the branching of the inflorescence showed phylogenetic significance and included dichasial, monochasial and unbranched patterns. In four of the East African Streptocarpus species sampled a single lateral bracteole was present on the first to third axes, after which the inflorescence was ebracteolate. Our results indicate that there may be some link between bracteole suppression and an alteration in the order of sepal initiation. The loss or suppression of lateral bracteoles also appears to result in the precocious development of the lateral cyme meristem.     

Growth and development in Arabidopsis thaliana through an entire life cycle under simulated microgravity conditions on a clinostat.

The effects of simulated microgravity conditions produced by a horizontal clinostat on the entire life cycle of Arabidopsis thaliana ecotype Columbia and Landsberg erecta were studied. Horizontal clinorotation affected little germination of seeds, growth and development of rosette leaves and roots during early vegetative growth stage, and the onset of the bolting of inflorescence axis and flower formation in reproductive growth stage, although it suppressed elongation of inflorescence axes. The clinorotation substantially reduced the numbers of siliques and seeds in Landsberg erecta, and completely inhibited seed production in Columbia. Seeds produced in Landsberg erecta on the clinostat were capable of germinating and developing rosette leaves normally on the ground. On the other hand, growth of pin formed mutant (pin/pin) of Arabidopsis ecotype Enkheim, which has a unique structure of inflorescence axis with no flower and extremely low levels of auxin polar transport activity, was inhibited and the seedlings frequently died during vegetative stage on the clinostat. Seed formation and inflorescence growth of the seedlings with normal shape (pin/+ or +/+) were also suppressed on the clinostat. These results suggest that the growth and development of Arabidopsis, especially in reproductive growth stage, is suppressed under simulated microgravity conditions on a clinostat. To complete the life cycle probably seems to be quite difficult, although it is possible in some ecotypes.     

Determination for inflorescence development is a stable state,separable from determination for flower development in Pisum sativum L. buds

Terminal meristems of Pisum sativum (garden pea) transit from vegetative to inflorescence development, and begin producing floral axillary meristems. Determination for inflorescence development was assessed by culturing excised buds and meristems. The first node of floral initiation (NFI) for bud expiants developing in culture and for adventitious shoots forming on cultured meristems was compared with the NFI of intact control buds. When terminal buds having eight leaf primordia were excised from plants of different ages (i.e., number of unfolded leaves) and cultured on 6-benzylaminopurine and kinetin-supplemented medium, the NFI was a function of the age of the source plant. By age 3, all terminal buds were determined for inflorescence development. Determination occurred at least eight nodes before the first axillary flower was initiated. Thus, the axillary meristems contributing to the inflorescence had not formed at the time the bud was explanted. Similar results were obtained for cultured axillary buds. In addition, meristems excised without leaf primordia from axillary buds three nodes above the cotyledons of age-3 plants gave rise to adventitious buds with an NFI of 8.3 ±0.3 nodes. In contrast seed-derived plants had an NFI of 16.5 ±0.2. Thus cells within the meristem were determined for inflorescence development. These findings indicate that determination for inflorescence development in P. sativum is a stable developmental state, separable from determination for flower development, and occurring prior to initiation of the inflorescence at the level of meristems.     

Evolutionary origin of the Asteraceae capitulum: Insights from Calyceraceae

? Premise of the study: Phylogenies based on molecular data are revealing that generalizations about complex morphological structures often obscure variation and developmental patterns important for understanding the evolution of forms, as is the case for inflorescence morphology within the well-supported MGCA clade (Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae). While the basal families share a basic thyrsic/thyrsoid structure of their inflorescences, Asteraceae possesses a capitulum that is widely interpreted as a racemose, condensed inflorescence. Elucidating the poorly known inflorescence structure of Calyceraceae, sister to Asteraceae, should help clarify how the Asteraceae capitulum evolved from thyrsic/thyrsoid inflorescences. ? Methods: The early development and structure of the inflorescence of eight species (five genera) of Calyceraceae were studied by SEM, and patterns of evolutionary change were interpreted via phylogenetic character mapping. ? Key results: The basic inflorescence structure of Calyceraceae is a cephalioid (a very condensed botryoid/thyrsoid). Optimization of inflorescence characters on a DNA sequence-derived tree suggests that the Asteraceae capitulum derives from a simple cephalioid through two morphological changes: loss of the terminal flower and suppression of the cymose branching pattern in the peripheral branches. ? Conclusions: Widely understood as a condensed raceme, the Asteraceae capitulum is the evolutionary result of a very reduced, condensed thyrsoid. Starting from that point, evolution worked separately only on the racemose developmental control/pattern within Asteraceae and mainly on the cymose developmental control/pattern within Calyceraceae, producing head-like inflorescences in both groups but with very different diversification potential. We also discuss possible remnants of the ancestral cephalioid structure in some Asteraceae.     

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.     

A novel gene Bractea (bra) controls the formation of an indeterminate bractless inflorescence in Arabidopsis thaliana

The morphological and genetic studies of the bra mutant of Arabidopsis thaliana (L.) Heynh. from the collection of the Department of Genetics and Breeding, Moscow State University, showed that the BRA gene controls the main stages of inflorescence development: it suppresses the development of leaflike organs subtending flowers (bracts) and inhibits the formation of the terminal flower. Inactivation of the BRA gene leads to the transition from the indeterminate bractless inflorescence characteristic of the family Cruciferaceae to the determinate bracteose inflorescence. The BRA gene plays a regulatory role and was probably involved in the conversion of the bracteose determinate inflorescence to the bractless indeterminate inflorescence during the origin of ancestral crucifers.