竹亚科簕竹属吊罗坭竹繁殖器官的补充描述

《中国植物志》第九卷第一分册及其它的相关文献均无关于竹亚科簕竹属吊罗坭竹花的描述或记录,本文作者采集制作了该竹子繁殖器官标本,并分别用中文和拉丁文对其进行了描述,其主要特征：花枝长70~170 cm;花序为总状花序或简单的圆锥花序;花序轴粗,节明显;小穗含4~5朵小花;小穗无柄;小穗轴节间无毛,呈棒状;颖2,膜质;外稃长或等于内稃;鳞被2;雄蕊6;子房卵形;柱头3,呈毛刷状;颖果未见。花期2~3月。花标本采自厦门市园林植物园。     

Floret Initiation and Development in Spring Wheat (Triticum aestivum L.)

The number and developmental stages of florets were determinedin each spikelet of the spike in the main shoots of spring wheat.Samples were taken frequently from plants grown in a phytotronand in a nitrogen application field-test. Ten stages of development,from floret initiation until anthesis, were recognized and described. Inter-spikelet variation in the total number of initiated floretswas rather small. However, the number of florets at advancedstages of development, as well as the number of grains, washighest in the central spikelets in which florets initiatedfirst. Floret initiation did not proceed beyond spike emergence,whereafter the distal florets and the spikelet apex degenerated.Grain-set was restricted to florets which had developed at leastto the stage of visible anther lobes at spike emergence. Thenumber of these florets was increased significantly by nitrogenapplication. Wheat, Triticum aestivum L., spikelet, floret, grain set, nitrogen     

糖蜜草（Melinis minutiflora Beauv．）幼穗分化发育．及花和果实形态的研究

本文对糖密草（ＭｅｌｉｎｉｓｍｉｎｕｔｉｆｌｏｒａＢｅａｕｖ．）的幼穗分化发育及花和果实的形态作了研究,将幼穗分化发育过程划分为以下九个时期：第一苞原基形成期;第一次枝梗原基形成期;第二、三次枝梗原基形成期;小穗及颖花原基形成期;雌、雄蕊原基形成期;花粉母细胞形成期;花粉母细胞减数分裂期;花粉充实期;花粉成熟期。全过程历时约需４２ｄ．从抽穗到颖果成熟约需５０ｄ。糖蜜草的花序为圆锥花序。每花序有可育花２０００—３０００朵．小穗是由小穗轴、内外颖片、不育花外稃和小花构成。小花包括有内外稃各一片、一鳞被、雄蕊三枚和一枚雌蕊,颖果千粒重为９１ｍｇ。     

Temperature Effects upon Inflorescence and Seed Development in Tall Fescue (Festuca arundinacea Schreb.)

The effects of three temperatures 15, 20, and 25 °C uponinflorescence and seed development in tall fescue (Festuca arundinaceaSchreb) between inflorescence emergence and seed maturity werestudied. Increasing temperature over this range reduced culmlength and the number of florets per spikelet, hastened theonset of anthesis and pollen release, increased relative growth-rateof the florets 9 days after peak anthesis, reduced the periodof seed development and 1000 seed weight No large effects oftemperature upon the percentage of florets setting seed werefound. The practical implications of these results are discussed.     

Heterochronic development of the floret meristem determines grain number per spikelet in diploid, tetraploid and hexaploid wheats

Background and Aims

The inflorescence of grass species such as wheat, rice and maize consists of a unique reproductive structure called the spikelet, which is comprised of one, a few, or several florets (individual flowers). When reproductive growth is initiated, the inflorescence meristem differentiates a spikelet meristem as a lateral branch; the spikelet meristem then produces a floret meristem as a lateral branch. Interestingly, in wheat, the number of fertile florets per spikelet is associated with ploidy level: one or two florets in diploid, two or three in tetraploid, and more than three in hexaploid wheats. The objective of this study was to identify the mechanisms that regulate the architecture of the inflorescence in wheat and its relationship to ploidy level.

Methods

The floral anatomy of diploid (Triticum monococcum), tetraploid (T. turgidum ssp. durum) and hexaploid (T. aestivum) wheat species were investigated by light and scanning electron microscopy to describe floret development and to clarify the timing of the initiation of the floret primordia. In situ hybridization analysis using Wknox1, a wheat knotted1 orthologue, was performed to determine the patterning of meristem formation in the inflorescence.

Key Results

The recessive natural mutation of tetraploid (T. turgidum ssp. turgidum) wheat, branching head (bh), which produces branched inflorescences, was used to demonstrate the utility of Wknox1 as a molecular marker for meristematic tissue. Then an analysis of Wknox1 expression was performed in diploid, tetraploid and hexaploid wheats and heterochronic development of the floret meristems was found among these wheat species.

Conclusions

It is shown that the difference in the number of floret primordia in diploid, tetraploid and hexaploid wheats is caused by the heterochronic initiation of floret meristem development from the spikelet meristem.Key words: Triticum, wheat, inflorescence, spikelet, floret, meristem, heterochrony, heterochronic development, knotted1, polyploidy     

Genetic control of branching patterns in grass inflorescences

Inflorescence branching in the grasses controls the number of florets and hence the number of seeds. Recent data on the underlying genetics come primarily from rice and maize, although new data are accumulating in other systems as well. This review focuses on a window in developmental time from the production of primary branches by the inflorescence meristem through to the production of glumes, which indicate the transition to producing a spikelet. Several major developmental regulatory modules appear to be conserved among most or all grasses. Placement and development of primary branches are controlled by conserved auxin regulatory genes. Subtending bracts are repressed by a network including TASSELSHEATH4, and axillary branch meristems are regulated largely by signaling centers that are adjacent to but not within the meristems themselves. Gradients of SQUAMOSA-PROMOTER BINDING-like and APETALA2-like proteins and their microRNA regulators extend along the inflorescence axis and the branches, governing the transition from production of branches to production of spikelets. The relative speed of this transition determines the extent of secondary and higher order branching. This inflorescence regulatory network is modified within individual species, particularly as regards formation of secondary branches. Differences between species are caused both by modifications of gene expression and regulators and by presence or absence of critical genes. The unified networks described here may provide tools for investigating orphan crops and grasses other than the well-studied maize and rice.

Recent work on grass inflorescence branching identifies extensive conserved regulation, but also divergence particularly in formation of secondary branches and spikelets.     

Interrelationship between Meristematic Regions of Developing Inflorescences of Four Cereal Species

A cultivar from each of four cereal species (Avena sativa L.,cv. Swan, Hordeum vulgare, L., cv. Clipper, Triticum aestivumL., cv. Gabo, and Secale cereale L., cv. South Australian Rye)was grown in controlled environment chambers in a 10-h photoperiod(short days) or 10-h photoperiod supplemented with a 6-h extensionby incandescent light. The developmental morphology of the inflorescenceswas followed to ascertain whether there were any common developmentalinterrelationships between the species. Inflorescence internodeelongation was initiated when the floret initial first appeared,irrespective of whether it occurred on the most advanced lateralspikelet or on the terminal spikelet of the rachis. The glumes(infertile bracts) of the terminal spikelet of the rachis wereinitiated when the first second-order inflorescence branch appeared,irrespective of whether the second-order inflorescence branchwas a floret initial or a lateral spikelet, as in Triticum sp.,or an inflorescence (panicle) branch, as in Avena sp. Cessationof the activity of the apical meristem, as measured by primordiumformation, was not correlated with any particular stage of floraldevelopment but appeared to be due to a lack of nutrients causedby an increasing competitiveness for the available nutrientsfrom the developing spikelets which are situated closer to thevascular system than the apical meristem.     

Studies on the Physiology of Flowering of Timothy (Phleum pratense L.) II. Influence of Daylength and Temperature on Size of the inflorescence

Inflorescence length in timothy increases when the photoperiodis reduced from 24 to 14 hours of light; it is also increasedby a reduction in ambient temperature from 75° to 55°F. There is a linear relation between total floret number andear length. Both factors affect ear length by influencing therate of growth of the spike between spikelet initiation andear emergence; this implies an effect on either the number ofprimary spikelet initials or the number of florets producedby branching, or both. Experiments with Lolium temulentum, wheredaylength and temperature influenced initiation and ear developmentin a way similar to that observed in timothy, suggest that thesefactors affect the number of florets at each primary initial.The interrelations of internal and external factors and theirinfluence on inflorescence size in the grasses is discussed.     

Comparative ontogeny of perfect and pistillate florets in Senecio vernalis (Asteraceae)

We studied the inflorescence, and in particular ontogeny and development of the florets in Senecio vernalis as a representative member of Asteraceae, using epi-illumination microscopy. Initiation and subsequent development of florets on the highly convex inflorescence apex occur acropetally, except for pistillate ray florets, which show a lag in initiation. Receptacular bracts derive from the receptacular surface after development of all florets. The order of whorl initiation in both disc and ray florets include corolla, androecium and finally the pappus, together with the gynoecium. Development of corolla lobes from a ring meristem occurs in bidirectional order starting from the lateral side, whereas stamens incept unidirectionally from the abaxial side. Concurrently with the inception of two median carpel primordia, a ring meristem develops at the base of the corolla from which pappus bristles differentiate in later stages. Pistillate ray florets show significant differences from perfect disc florets as reflected by the zygomorphic shape of the floral apex and a shift of floral merosity from pentamery to tetramery. Loss of stamens in ray florets occurs due to abortion of primordia after initiation.     

ABERRANT PANICLE ORGANIZATION 1 temporally regulates meristem identity in rice

We report a recessive mutation of rice, aberrant panicle organization 1 (apo1), which severely affects inflorescence architecture, floral organ identity, and leaf production rate. In the wild-type inflorescence, the main-axis meristem aborts after forming 10-12 primary branch primordia. However, in apo1, the main-axis meristem was converted to a spikelet meristem after producing a small number of branch primordia. In addition, the branch meristems in apo1 became spikelet meristems earlier than in wild type. Therefore, in the inflorescence, the apo1 mutation caused the precocious conversion of the meristem identity. In the apo1 flower, lodicules were increased at the expense of stamens, and carpels were formed indeterminately by the loss of meristem determinacy. Vegetative development is also affected in the apo1. Leaves were formed rapidly throughout the vegetative phase, indicating that APO1 is also involved in temporal regulation of leaf production. These phenotypes suggest that the APO1 plays an important role in the temporal regulation of both vegetative and reproductive development.     

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.     

Effects of shading on inflorescence size and development in temperate perennial grasses

Single plants of S24 perennial ryegrass (Lolium perenne L.) and S215 meadow fescue (Festuca pratensis Huds.) were transferred between, or exposed continuously to, contrasting light intensities obtained by decreasing the natural light in a glasshouse with Tygan shades. Inflorescence development in main shoots was studied by dissections of shoot apices, and by counts of branches and florets when ears emerged. Apical growth was slower, and spikelet initiation and inflorescence development were delayed or inhibited, in decreased light intensities. The number of main branches in the ear depended on the rate of apical growth before and after spikelet initiation, and on the time of spikelet initiation. In meadow fescue these processes were influenced by light intensity. Floret numbers per inflorescence branch were generally decreased by decreased light intensity. Most of the effects of light intensity on inflorescence development were smaller in ryegrass than in meadow fescue.     

DEVELOPMENTAL STUDIES IN PTYCHOSPERMA (PALMAE). I. THE INFLORESCENCE AND THE FLOWER CLUSTER

This paper describes inflorescence structure, including organogenesis of the panicle and flower clusters and vasculature of flowering branches, for two species of Ptychosperma, a genus of arecoid palms. The inflorescence is an infrafoliar panicle with up to four orders of branches in a spirodistichous arrangement conforming to an irregular one-half phyllotaxy. The primordium of the inflorescence is crescentic and the apex has two tunica layers, a group of central cells, and a rib meristem. The distal flower-bearing parts or rachillae of all branches develop acropetally early in ontogeny and are vertically oriented in the bud. Although these rachillae terminate branches of different sizes and orders, they are similar in size and in number of flower clusters produced. Internodes and lower parts of branches develop later. Bracts of four types are produced: a prophyll and empty peduncular bract, bracts which subtend lateral branches, bracts subtending triads, and floral bracteoles. The prophyll and peduncular bracts are tubular and completely closed around all branches until about three months before the flowers reach anthesis. Bracts subtending lateral branches and those that subtend triads enlarge by small amounts of apical, adaxial, and marginal growth to cover subtended apices during early ontogeny, but are small to absent at maturity. Flower clusters are triads of two lateral staminate and a central pistillate flower. Organogenesis indicates that the triad is a sympodial unit. Flowers develop successively, each floral apex bearing a bracteole that subtends the next flower. The vasculature of the inflorescence may be divided into two systems. Bundles of the main axis extend acropetally into the vertically oriented branches as they are initiated and form a central cylinder of larger bundles in each branch. Flower clusters are supplied by a peripheral system of smaller bundles that develop later in relation to the developing floral organs. Bundles of the peripheral system branch frequently, but branching levels are irregular. The irregular branching of peripheral bundles appears related to the phyllotaxy of the flower clusters and the random right or left position of the first flower of the triad. The level of branching of a bundle may depend on the position of a floral primordium with respect to an existing procambial strand. Three (-4) bundles supply each staminate flower and six (-10) the pistillate flower. The histologically specialized inflorescence has stomata and contains abundant starch. Tannins and raphides, spherical silica bodies, and various forms of sclerenchyma appear in sequence and apparently provide support and protection during the long exposure of the branches.     

Reproductive morphology of the early-divergent grass Streptochaeta and its bearing on the homologies of the grass spikelet

Reproductive morphology and development are described in the Brazilian grass Streptochaeta spicata, in order to assess the homologies of the characteristic grass inflorescence, termed a spikelet, and other reproductive organs. Streptochaeta possesses some features that are commonly found in Poaceae, including a well-differentiated embryo. It also possesses some relatively unusual, presumably derived features, such as non-plumose stigmas, which indicate that it could be insect-pollinated. It shares some features with other early-divergent grasses, such as Pharus, which could represent plesiomorphic conditions for grasses. The inflorescence unit in Streptochaeta has been interpreted as a compound branching system or pseudospikelet. The present data suggest that it is a highly modified spikelet, with a modified flower borne either on a different axis to the basal bracts (glumes) or on the same axis as the basal bracts. The three bracts below the stamens are interpreted as homologous to the lodicules. The Streptochaeta spikelet could be considered as morphologically intermediate between the true spikelet of grasses and reproductive units of close grass relatives.     

BRANCHED SILKLESS mediates the transition from spikelet to floral meristem during Zea mays ear development

The molecular and genetic control of inflorescence and flower development has been studied in great detail in model dicotyledonous plants such as Arabidopsis and Antirrhinum . In contrast, little is known about these important developmental steps in monocotyledonous species. Here we report the analysis of the Zea mays mutant branched silkless1–2 (bd1–2) , allelic to bd1 , which we have used as a tool to study the transition from spikelet to floret development in maize. Floret development is blocked in the female inflorescence (the ear) of bd1–2 plants, whereas florets develop almost normally in the male inflorescence (the tassel). Detailed phenotypic analyses indicate that in bd1–2 mutants ear inflorescence formation initiates normally, however, the spikelet meristems do not proceed to form floret meristems. The ear spikelets, at anthesis, contain various numbers of spikelet-like meristems and glume-like structures. Furthermore, growth of branches from the base of the ear is often observed. Expression analyses show that the floral-specific MADS box genes Zea mays AGAMOUS1 ( ZAG1 ), ZAG2 and Zea mays MADS 2 ( ZMM2 ) are not expressed in ear florets in bd1–2 mutants, whereas their expression in tassel florets is similar to that of wild type. Taken together, these data indicate that the development from spikelet to floret meristem is differentially controlled in the ear and tassel in the monoecious grass species Zea mays , and that BRANCHED SILKLESS plays an important role in regulating the transition from spikelet meristem to floral meristem during the development of the female inflorescence of maize.