Reproductive Development of the Apex of Brachiaria decumbens Stapf

The development of Brachiaria decumbens tillers, as based onapex morphology, may be conveniently divided into six phases.These are the (1) vegetative, (2) raceme initiation, (3) spikeletinitiation, (4) spikelet differentiation, (5) inflorescenceexsertion and anthesis, and (6) seed maturation phases. Theonset of reproductive development is characterized by an increasein apex length and proceeds with the expansion of a bud in theaxil of the most recently initiated leaf primordium. This budgives rise to the first raceme and further racemes are formedin basipetal succession. Changes in apex morphology during thefirst four phases of development are described and illustratedwith scanning electron microscope and median longitudinal sectionphotomicrographs. Brachiaria decumbens, signal grass, apex morphology, SEM, median longitudinal sections, developmental phases     

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.     

Physiology of Flowering in Saccharum: I. DAYLENGTH CONTROL OF FLORAL INITIATION AND DEVELOPMENT IN S. SPONTANEUM L.

Flowering in two clones of Saccharum spontaneum L. is controlledby photoperiod. The earliest stages of development, ‘induction’and ‘initiation of the inflorescence axis primordium’(IAP) were optimally promoted under intermediate days of 12h 30 min, while the subsequent stage ‘initiation of inflorescencebranch primordia’ (IBP) was inhibited by days longer than13 h. The following stage ‘initiation of spikelet primordia’(ISP) showed a quantitatively intermediate response with anoptimum photoperiod of 9 h to 11 h. The elongation of the differentiatedinflorescence was found to be only slightly sensitive to photoperiodsof 13 h or longer in one of the clones. Unfavourable photoperiodsat stages following induction resulted in the arrest or delayof inflorescence development and when these were given duringthe IAP and IBP stages, reversion to the vegetative conditioncommonly occurred.     

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     

The presentation of treatment responses from block experiments after analysis of variance of transformed data

In pearl millet, severe water deficit during the period of panicle development delays flowering. The flowering response of both main shoot and tillers to water stress during panicle development was investigated using four hybrids. Panicle initiation of all tillers occurred in the three early genotypes despite water stress. In the late genotype, however, panicle initiation of tillers occurred only after the release of stress. The delay in flowering due to water stress was more pronounced in the tillers than in the main shoot. However, the proportion of tillers producing an inflorescence was increased by water stress. Grain yield losses on the main shoot by water stress were compensated by an increase in tiller grain yields. Delay in flowering and buffering by tillers provide an important adaptive mechanism to overcome a period of drought stress prior to flowering.     

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.     

Density Studies on Lupins: I. Flower Development

In an August-sown experiment the pattern of flower developmentwas followed for cv. Ultra (Lupinus albus L.) and cv. Unicrop(L. angustifolius L.) grown at low (10 plants m^–2) andhigh (93 and 83 plants m^–2, Ultra and Unicrop respectively)densities. Dry weight increase of flowers on the main-stem inflorescenceand first lateral below the main-stem were compared at differentfloral stages. Maximum flower weight was reached just priorto the open flower stage and remained constant or declined untila pod formed or abscission occurred. The time period betweenmaximum flower weight and pod formation or abscission was upto 10 days. Emergence of the inflorescence was earlier and thefirst flower of Ultra opened 10 days before Unicrop. Developmentof each terminal raceme (inflorescence) was acropetal, withpods having formed on lower flower nodes when terminal flowerswere still quite immature. Laterals forming the next generationof inflorescences grew from axillary leaf buds below an inflorescencewhile it was in full flower. Sources of competition from connectedreproductive and vegetative metabolic sinks are discussed. Lupinus spp., lupins, flower development, planting density     

Tiller dynamics and assimilate partitioning in Lolium perenne with particular reference to flowering

The fate of 100 seedling plants of Lolium perenne L. was studied over a period of 2 years in a field plot. The birth and death of tillers and the production of inflorescences was followed, and the components of seed yield were recorded in detail in the first year. The pattern of distribution of ¹⁴CO₂ assimilated by the main shoot was examined at monthly intervals and during the first flowering season the distribution of ¹⁴C-assimilate from individual leaves and from the inflorescence was also studied. The capacity of individual tillers to assimilate ¹⁴CO₂ prior to flowering and the re-distribution of previously accumulated assimilate during seed growth were also assessed. Plants died at a more or less constant rate with time and only 54 survived to the end of the 2–yr period. First year mortality was associated with severe grazing or cutting but in the second year the death of ungrazed plants was observed. There was great variability in the production of tillers by surviving plants. In both years the number of live tillers per plant increased from July to the end of April with particularly rapid tillering in March and April establishing the maximum value for each year. There was a similar phase of rapid tillering after flowering in July. The number of live tillers per plant declined by 50% during stem elongation and inflorescence emergence and the majority of dead tillers were young secondary (in the first year) and tertiary (in the second year) tillers with a mean age of 40 days. Such tillers had poor assimilatory capacity prior to the onset of death and were not supplied with assimilate from the main shoot. Most of the plants surviving at the end of the experiment flowered in both years and one quarter of the maximum number of live tillers per plant recorded in April of each year produced inflorescences. The earlier a tiller was produced the greater was its chance of flowering and the greater its production of seed. The greater weight of seed produced was associated with the development of more seed-bearing florets per spikelet. There was relatively little export of “C-assimilate from the flowering main shoot, and the lower internodes formed the major sink for post-anthesis assimilate. The growth of seeds appeared to be relatively independent of the leaves for current assimilate. There was some evidence that assimilate accumulated in lower internodes was remobilised and utilised in the growth of seeds and new tillers. Overall, the results confirm the view that the grass plant is a dynamic population of short-lived tillers and indicate that increasing competition for assimilate at flowering exerts a major influence on the production and survival of tillers.     

野生翠雀传粉生物学特性研究

对从北京郊区引种的野生翠雀(Delphiniun grandiflorum L.)的开花习性进行了观察,对其花粉活力、柱头可授性,花粉/胚珠比(P/O),杂交指数(OCI)进行了检测,并进行了人工授粉实验。结果表明:(1)翠雀的花序为总状花序,具3~15朵花,花梗长1.5~3.8cm,单花期10d,花序花期15~18d。(2)翠雀花粉活力日变化呈先升高后降低的趋势;柱头在开花当天至开花后4d无可授性,在开花后的5d、6d有可授性。(3)翠雀的花粉胚珠比(P/O)为2 159.14,杂交指数OCI为5;人工授粉实验表明,翠雀在自然条件下能够结实,结实率为100%;存在自主自花授粉现象,部分自交亲和,结实受传粉者限制;不存在无融合生殖现象。(4)翠雀主要的访花昆虫有四条无垫蜂和短腹管蚜蝇。研究认为,翠雀的繁育系统为兼性异交或专性异交,部分自交亲和,需要传粉者。     

The Effects of Daylength and Treatment with Gibberellic Acid on Spikelet Initiation and Development in Clipper Barley

The effects of photoperiod, light quality and a single applicationof gibberellic acid (GA₃) on the development of the main-stemapex in Clipper barley are reported. In 16 and 24 h days spikeletinitiation was rapid but extended over a short period whereasin 8 h photoperiods both spikelet initiation and developmentwere slower but occurred for a much longer time. Initiationalways stopped when the anther primordia were clearly visiblein the most advanced spikelet. Daylength extensions with lowintensity incandescent light were most effective when they followedrather than preceeded the 8 h period of high light intensity.Plants grown in 8 h high intensity followed by 8 h low intensitylight initiated spikelets almost as rapidly as those grown in16 h high intensity light. Thus, the effects of daylength onspikelet production were primarily mediated through photoperiodicallycontrolled processes rather than through photosynthesis andassimilate supply. The effects of applied GA₃ were long livedand greatest in short days where the rates of both spikeletinitiation and development were promoted. The parallels betweenthe effects of long days and GA₃ treatment are discussed togetherwith possible reasons for the cessation of initiation and thelong duration of the GA₃ effect. daylength, gibberellic acid, spikelet initiation, Hordeum vulgare L., barley, main-stem apex, primordia     

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.     

SPIRAL DEVELOPMENTAL PATTERNS IN THE STEM AND INFLORESCENCE OF LILIUM TIGRINUM

Extensive correlations in spirality were observed among vegetative and floral organs in Lilium tigrinum Ker. Organs involved were vegetative leaves, bracts, and bracteoles. These correlations varied in their degree of constancy depending upon the organs involved. The mature inflorescences of L. tigrinum appeared to fit the common definition of a raceme. In 67.3% of the flowers at node 3 on the raceme, the bract-bracteole spirals reversed the spiral of vegetative leaves on the stem. These reversals resembled those observed on essentially cymose inflorescences of certain members of the Caryophyllaceae. Cymose branching was found to be an invariable feature of the inflorescence of L. tigrinum when secondary flowers appear. The apparently indeterminate tips of inflorescence main axes were interpreted as exhibiting stages in progression from a basically determinate (cymose) inflorescence. It was concluded that the ancestors of L. tigrinum had well-developed cymose branching patterns in the inflorescence. Reversal of stem spirals by the bract-bracteole spirals at the apices of many inflorescences was considered to be the result of complete utilization of the inflorescence meristem. Explanations for those reversals were provided by the field theory and by the theory of the first available space.     

Characterization and mapping of a Prbs gene controlling spike development in Hordeum vulgare L.

The barley mutant, poly-row-and-branched spike (prbs) showed altered inflorescence morphology: complete conversion of the rudimentary lateral spikelets in two-rowed barley into fully developed fertile spikelets similar to the six-rowed phenotype, and additional spikelets in the middle of spike. Moreover, branched spikes emerged in progeny from a cross between the mutant and a six-rowed barley cultivar. Morphological observation of the development of immature spikes of the mutant and descendants with branched spikes showed that the Prbs gene is involved in spikelet development in the triple-mound stage. In mutant prbs, new meristems initiated at the flanks of lateral spikelets and middle spikelet meristems were converted to branch meristems, developing branched spikes. These observations suggested that the Prbs gene plays a crucial role in spikelet initiation and identity maintenance. The Prbs gene may be an important modifier in inflorescence differentiation from a panicle into a spike. The branched spikes emerging in hybrids from a cross between the mutant and six-rowed barley cultivar were not conferred by the gene vrs1 or Int-c, which decide spike morphology in six-rowed barley. These results imply that although six-row genes vrs1 and Int-c and prbs have similar effects on lateral spikelet development, they have different functions in branched spikes. The Prbs gene was mapped to chromosome 3H between SSR marker Bmag0023 and marker Cbic60 at a genetic distance of 3.3 and 5.4 centimorgans (cM), respectively.     

Callus initiation and plant regeneration from inflorescence primordia of the intergeneric hybrid Agropyron repens (L.) Beauv.xBromus inermis Leyss. cv. nanus on a modified nutritive medium

Plant regeneration from callus of intergeneric hybrid Agropyron repens (L.) Beauv. x Bromus inermis Leyss cv. nanus (AGROMUS) was carried out on a new culture medium designated medium-F. Within 21 days of the plating of inflorescence primordia the initiated callus showed globular structures. From the 21st day of culture, one step plant regeneration occurred on the callus without subculture. The new basal medium reported in this work was effective in callus initiation and plant regeneration of the hybrid AGROMUS by (i) the reduction of the total ion strength (2.6 g/l, 22.5 mM) of macroelements compared to MS (4.5 g/l,45.2 mM), (ii) the use of NH₄NO₃ as the sole N-source, and (iii) the application of KH₂PO₄ at an 8 times higher concentration (1160 mg/l,8.5 mM) when compared to the Murashige and Skoog medium composition. This medium provided a 2 to 10 fold reduction in the 2,4-dichlorophenoxyacetic acid supplement needed for the callus initiation and one step plant regeneration after a gibberellic acid (2 mg/l, for 5 days) pretreatment of tillers. The regenerated plantlets were subcultured in multi-shoot culture and potted in soil to grow for further analysis.Abbreviations AA amino acid medium (Müller and Grafe 1978, Toriyama and Hinata 1985) - B5 Gamborg et al. (1968) medium - 2,4-D 2,4-dichlorophenoxyacetic acid - GA₃ gibberellic acid - MS Murashige and Skoog (1962) medium - NAA naphthaleneacetic acid - N6 Chu et al. (1975) medium - SH Schenk and Hildebrandt (1972) medium - 2,4,5-T 2,4,5-trichlorophenoxyacetic acid     

Effects of temperature and nitrogen supply on the growth of timothy (Phleutn pratense L.)

Timothy was grown in controlled environments with day/night temperatures of 18.5/10° C. and 29.5/21° C, with and without extra nitrogen. High temperatures hastened all stages of development after inflorescence initiation. Anthesis occurred after 48 days when N was added and after 52 days without N. Anthesis did not occur until after 68 days at the lower temperatures, with or without N. Largest yields of leaves, stems and stubble were produced at all growth stages at the lower temperatures with N. This treatment gave the highest crop growth rate, which increased almost linearly with time. The lower temperatures with N gave many more and longer stems and more inflorescences than any other treatment. With N and high temperatures, there was no increase in crop growth rate after inflorescence emergence. Temperature had little effect on growth when no N was provided. With no added N, crop growth rate was retarded and increased very little after inflorescence initiation. Herbage produced at high temperatures had the lower percentages of water-soluble carbohydrates and digestible dry matter, but had the higher percentages of most minerals. N increased the percentage content of most minerals. Weights of stubble and of water-soluble carbohydrates in the stubble, and the numbers of stem bases and tillers, were larger at the lower temperatures, and were usually larger with added N in both temperature regimes. Nitrogen decreased the percentage content of carbohydrates in the stubble. Large haplocorms were produced at the low temperatures, but none developed at the high temperatures, indicating less carbohydrate storage. The results partly explain the vigorous growth of timothy in the cool, temperate regions of Europe and North America, and the small dry matter production, lower nutritional value of the herbage, and lack of persistence in the warm southern regions where rainfall is adequate.     

MORPHOLOGY,INHERITANCE, AND EVOLUTIONARY SIGNIFICANCE OF SEX REVERSAL IN TRIPSACUM DACTYLOIDES (POACEAE)

Inflorescences of Tripsacum dactyloides (L.) L. (Andropogoneae) are characterized by single female spikelets at one to several basal nodes and paired male spikelets at several nodes above them on each raceme. Female spikelets are one-flowered and male spikelets are two-flowered. A sex form variant was found in a wild population in north central Kansas and classified as T. dactyloides (L.) L. forma prolificum Dayton et Dewald. The variant of this native distant relative of maize (Zea mays L. spp. mays) differs from the normal form by having both pistillate and perfect rather than staminate spikelets in the terminal (tassel) portion of the inflorescence and by having two functional pistillate florets in the basal spikelets instead of one. A recessive major gene at a single locus regulates the change of the inflorescence from monoecious to gynomonoecious.     

Size of the Chrysanthemum Shoot Apex in Relation to Inflorescence Initiation and Development

The size of the apical dome of Chrysanthemum morifolium Ramat.at the transition to inflorescence initiation in continuouslight (long days) was not systematically influenced by eitherthe temperature or the irradiance under which the plants weregrown. It was generally 0.26 mm in diameter and c. 3.6 x 10^–3mm³ in volume when the first bract was initiated. The dimensionsof the apical dome of plants in short days were only slightlysmaller at this stage. Similarly, each step in the further developmentof the chrysanthemum inflorescence was associated with a narrowrange of apex sizes, indicating that inflorescence initiationand development are closely related to apex size. Chrysanthemum morifolium Ramat, shoot apex, inflorescence initiation     

ESSENTIAL OILS AS TAXONOMIC AIDS IN THE CLASSIFICATION OF DICHANTHIUM PARVIFLORUM

Essential oil components and gross morphological characters are closely correlated in Dichanthium parviflorum (R. Br.) de Wet et Harlan (Gramineae) and related species. Different species, varieties, and geographical races, as well as hybrids between them, can be identified on the basis of absence or presence and quantity of essential oil components. The morphologically variable D. parviflorum was subdivided into four varieties: var. parviflorum, var. capilliflorum (Steud.) de Wet et Harlan comb. nov., var. mutispiculum (Ohwi) de Wet et Harlan comb. nov., and var. spicigerum (S. T. Blake) de Wet et Harlan comb. nov. These varieties differ from each other morphologically in having respectively racemes with 1-4 and awned, 3-5 and awned, 1-2 and awnlass, and 4-10 and awned spikelet pairs per raceme.