The CUP-SHAPED COTYLEDON2 and 3 genes have a post-meristematic effect on Arabidopsis thaliana phyllotaxis

Background and Aims The arrangement of flowers in inflorescence shoots of Arabidopsis thaliana represents a regular spiral Fibonacci phyllotaxis. However, in the cuc2 cuc3 double mutant, flower pedicels are fused to the inflorescence stem, and phyllotaxis is aberrant in the mature shoot regions. This study examined the causes of this altered development, and in particular whether the mutant phenotype is a consequence of defects at the shoot apex, or whether post-meristematic events are involved.Methods The distribution of flower pedicels and vascular traces was examined in cross-sections of mature shoots; sequential replicas were used to investigate the phyllotaxis and geometry of shoot apices, and growth of the young stem surface. The expression pattern of CUC3 was analysed by examining its promoter activity.Key Results Phyllotaxis irregularity in the cuc2 cuc3 double mutant arises during the post-meristematic phase of shoot development. In particular, growth and cell divisions in nodes of the elongating stem are not restricted in the mutant, resulting in pedicel–stem fusion. On the other hand, phyllotaxis in the mutant shoot apex is nearly as regular as that of the wild type. Vascular phyllotaxis, generated almost simultaneously with the phyllotaxis at the apex, is also much more regular than pedicel phyllotaxis. The most apparent phenotype of the mutant apices is a higher number of contact parastichies. This phenotype is associated with increased meristem size, decreased angular width of primordia and a shorter plastochron. In addition, the appearance of a sharp and deep crease, a characteristic shape of the adaxial primordium boundary, is slightly delayed and reduced in the mutant shoot apices.Conclusions The cuc2 cuc3 double mutant displays irregular phyllotaxis in the mature shoot but not in the shoot apex, thus showing a post-meristematic effect of the mutations on phyllotaxis. The main cause of this effect is the formation of pedicel–stem fusions, leading to an alteration of the axial positioning of flowers. Phyllotaxis based on the position of vascular flower traces suggests an additional mechanism of post-meristematic phyllotaxis alteration. Higher density of flower primordia may be involved in the post-meristematic effect on phyllotaxis, whereas delayed crease formation may be involved in the fusion phenotype. Promoter activity of CUC3 is consistent with its post-meristematic role in phyllotaxis.     

On the diffusion theory of phyllotaxis

An inhibitor diffusion theory of phyllotaxis is examined in the steady-state approximation for cylindrical shoot apex models. The model calculations give rise naturally to common patterns of spiral phyllotaxis, as well as to higher whorled patterns. The model also predicts commonly observed subpatterns of axillary organs superimposed on primary phyllotaxis patterns. Application of the model to phyllotaxis patterns in other organisms and in flowers is proposed.     

The Shoot Apex and its Dichotomous Branching in the Nypa Palm

Seedlings of the palm Nypa fruticans van Wurmb are viviparous,the plumule becoming exserted before the fruit is ripe and possiblyassisting in fruit detachment. Established seedlings have horizontalaxes, this growth orientation being maintained throughout thelife of the palm which may be described as ‘rhizomatous’.Inflorescences are axillary in adult plants, but their distributionis irregular. The shoot apex is small and very asymmetricalsince it is more or less displaced into a lateral position byprogressive enlargement of each leaf primordium during thisleaf's first plastochrone. The plastochrone interval is apparentlya long one so that leaves have a wide developmental gap betweenthem. This results in a leaf base which is more or less cylindricalbut with a groove to accommodate the next youngest leaf. All the available evidence suggests that vegetative branchingis by equal dichotomy of the shoot apex at wide intervals. Thedichotomy is marked superficially by a leaf, enclosing the twonew shoots, which has two grooves. The twin shoots are insertedin the lateral, not the dorsiventral plane of this enclosingleaf. The daughter shoots are always at precisely the same stageof development and they always have mirror-image symmetry withrelation to each other, the phyllotaxis of the parent shootbeing maintained without obvious interruption. There is no anatomicalevidence for abortion of the original apex and its replacementby two new ones. The vascular system is divided equally betweeneach daughter shoot without interruption, suggesting stronglythat there is continuity of growth from the undivided to thedivided condition.     

Ontogeny of the reproductive shoot apex ofClethra barbinervis,especially on the superficial view

The structure and the ontogenetic process of the reproductive shoot apex forming a terminal inflorescence ofClethra barbinervis were examined, especially concerning the superficial view of the apex. The system of contact parastichies is 2+3 in phyllotaxis in the vegetative phase, changing to 5+8 for bract arrangement in the reproductive phase. At the same time the size of the apex is conspicuously enlarged. The size of the foliage leaf primordia in the vegetative phase is larger than that of the bract primordia in the reproductive phase. The radial cell files, which are clear in the vegetative shoot apex, are not recognizable at least in the early stage of the reproductive phase. The author proposes a close correlation between the appearance of the radial cell files, as well as the construction of the apical sectors, and the sizes of the shoot apex and leaf primordia. It may be proposed also that the construction of the apical sectors is closely correlated with the phyllotaxis.     

Modeling of Branching Patterns in Plants

A major determinant of plant architecture is the arrangement of branches around the stem, known as phyllotaxis. However, the specific form of branching conditions is not known. Here we discuss this question and suggest a branching model which seems to be in agreement with biological observations. Recently, a number of models connected with the genetic network or molecular biology regulation of the processes of pattern formation appeared. Most of these models consider the plant hormone, auxin, transport and distribution in the apical meristem as the main factors for pattern formation and phyllotaxis. However, all these models do not take into consideration the whole plant morphogenesis, concentrating on the events in the shoot or root apex. On the other hand, other approaches for modeling phyllotaxis, where the whole plant is considered, usually are mostly phenomenological, and due to it, do not describe the details of plant growth and branching mechanism. In this work, we develop a mathematical model and study pattern formation of the whole, though simplified, plant organism where the main physiological factors of plant growth and development are taken into consideration. We model a growing plant as a system of intervals, which we will consider as branches. We assume that the number and location of the branches are not given a priori, but appear and grow according to certain rules, elucidated by the application of mathematical modeling. Four variables are included in our model: concentrations of the plant hormones auxin and cytokinin, proliferation and growth factor, and nutrients—we observe a wide variety of plant forms and study more specifically the involvement of each variable in the branching process. Analysis of the numerical simulations shows that the process of pattern formation in plants depends on the interaction of all these variables. While concentrations of auxin and cytokinin determine the appearance of a new bud, its growth is determined by the concentrations of nutrients and proliferation factors. Possible mechanisms of apical domination in the frame of our model are discussed.     

Formation of Pseudowhorls inPeperomia verticillata(L.) A. Dietr. Shoots Exhibiting Various Phyllotactic Patterns

Pseudowhorls are composed of leaves attached at almost equallevels and separated by single fully elongated internodes. InPeperomiaverticillata, pseudowhorls form regularly in shoots exhibitingboth spiral and truly whorled patterns of phyllotaxis. In spiralsystems, they are composed of successive leaves positioned onthe ontogenetic helix. In whorled phyllotaxis, leaves of twoadjacent whorls occur at almost the same level and this wayform a pseudowhorl. The number of leaves per pseudowhorl dependson the type of phyllotactic pattern and also the system of primordiapacking. In all the shoots, regardless of the type of phyllotaxis,the number of leaves per pseudowhorl equals the number of leafprimordia in physical contact with the apical dome. It is thesame as the higher number in contact parastichy pairs in spiralpatterns or the number of orthostichies in whorled phyllotaxis.The pseudowhorled pattern is already manifested in the arrangementof leaf primordia. In spiral and whorled phyllotaxis the plastochronratio calculated for primordia or whorls belonging to adjacentpseudowhorls is always higher than that calculated for membersof one pseudowhorl. Moreover, angular distances between primordiaof one pseudowhorl in spiral patterns are more uniform thanexpected in Fibonacci phyllotaxis. These observations were madeon plants both growing in pots and culturedin vitro. 6-Benzylaminopurine,a synthetic cytokinin, added to the medium increases the meannumber of leaves per pseudowhorl. It seems that this effectis indirect: phyllotaxis changes first rather than the destinyof a particular internode in a process of selective elongation.Copyright1999 Annals of Botany Company Peperomia verticillata, pseudowhorls, phyllotaxis, shoot apex.     

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.     

Phyllotaxis, phenology and architecture in Cephalotaxus, Torreya and c(Coniferales)

In Amentotaxus, Cephalotaxus and Torreya there is a regular seasonal alternation of foliage leaves and bud-scales, with foliage leaves largely preformed, i.e. initiated in the season before they expand. On most plagiotropic shoots phyllotaxis in the production of foliage leaves may be either bijugate ( Cephulotaxus, Torreya ) or decussate ( Amentotaxus ). In bijugate phyllotaxis successive leaf pairs originate at an angle of about 68° to each other, i.e. approximately one-half of the 'ideal' or Fibonacci angle of 137.5°. Secondary leaf orientation in Cephulotaxus and Torreya , by twisting of the leaf base, produces the dorsiventrality of plagiotropic shoots, whereas in Amentotaxus secondary orientation involves a twisting of the stemc as well as the leaf base. In Cephalotaxus cc condition is constant in the production of the numerous but imprecise number of bud-scales and in the production of foliage leaves. However, in Torreya the phyllotaxis changes from bijugate in the production of foliage leaves to decussate in the production of bud-scales, which are constant in number (about eight pairs). This allows a precise analysis of the biphasic production of leaf primordia in the seasonal cycle. The phyllotactic change in Torreya may not be the result of reported changes in shoot apex dimensions since Cephalotaxus , with its constant phyllotaxis, has a comparable seasonal change in apex dimensions. Information on architecture, chirality and cone morphology is also included.     

Experimental and Analytical Studies of Pteridophytes: XXXVIII. Some Observations on Spiral and Bijugate Phyllotaxis in Dryopteris aristata Druce

In an attempt to assess some of the factors which determineleaf arrangement at the shoot apex, a study has been made ofphyllotaxis in the fern Dryopteris aristata. It is shown thatclockwise and counter-clockwise spirals occur with equal frequencyboth in field collections and in adventitious and experimentallyinduced buds which arise on the rhizomes and apices of plantsgrown in the laboratory. In addition to the two common spiralarrangements, a form with bijugate phyllotaxis has been noted.It is concluded that the three types of leaf arrangement arenot under direct genetic control, but rather are dependent oncircumstances of growth at the apex.     

灌木铁线莲（毛茛科）花器官的发生与发育

用扫描电子显微镜（SEM）对铁线莲属（Clematis L.）植物灌木铁线莲(C. fruticosa Turcz.)花的形态发生和发育过程进行了观察。灌木铁线莲花原基形成后,4枚萼片以交互对生的方式首先发生,呈轮状排列。最早的4枚雄蕊原基在4枚萼片交接的位置上近螺旋状发生,此后,随着雄蕊原基的向心发生和数目不断增多,其发生的螺旋状序列逐渐明显。雄蕊原基发生后,在花原基顶端,心皮原基沿着雄蕊原基的发生序列呈螺旋状发生。本文结果支持在原始被子植物花中螺旋状排列和轮状排列同时存在的观点。此外,本文也进一步证实了花萼与苞片的同源性。     

Growth changes in the shoot apex of Sinapis alba during transition to flowering

The aim of the work was to report morphological changes whichoccur in the shoot apex during the morphogenetic switch to floweringin the model long day (LD) plant, Sinapis alba. During the floraltransition induced by 1 LD the growth rate of all componentsof the shoot apex is modified profoundly. The earliest changes,detected at 24 h after start of LD, include a decrease in plastochronduration and an increase of growth of leaf primordia. One daylater, the meristem dome starts to increase in volume, apicalinternodes have an increased height and there is a precociousoutgrowth of axillary meristems. All these changes precede initiationof flower primordia, which starts at about 60 h after the startof LD. Later changes include meristem doming, a decrease inthe plastochron ratio and a shift to a more complex phyllotaxis.All the changes, except the decreased plastochron ratio, arecharacteristics of an apex with an increased tempo of growth.The stimulation of longitudinal growth (height of apical intemodes)is more marked and occurs earlier than the reduction of radialgrowth (plastochron ratio). Key words: Axillary meristem, internode growth, leaf growth, plastochron ratio, plastochron duration     

Phytoplasma distribution in coconut palms affected by lethal yellowing disease

Lethal yellowing (LY), the most devastating disease affecting the coconut palm in America, is caused by phytoplasmas known to be distributed in different parts of infected plants. However, no comprehensive reports exist on the phytoplasma distribution. This study refers to the detection of LY phytoplasma DNA using PCR in different coconut plant parts, throughout the development of the disease. Sample analysis of positive palms taken at different stages of disease development (either symptomatic or symptomless) showed differences in the percentage of LY detection between plant parts. Some parts showed a very high level of LY DNA (stem, young leaves, inflorescences, stem apex and root apex), low levels were found in the intermediate leaves and roots without apex, whereas no LY phytoplasma DNA was detected in mature leaves. The detection percentage of LY phytoplasma DNA was lowest in symptomless‐infected palms for all parts, except the stem, where phytoplasma accumulations were consistently detected. This pattern of detection among parts is consistent with the hypothesis that phytoplasmas move from photosynthate source tissues to sink tissues via the phloem mass flow process. The accumulations in the (lower) stem, prior to the appearance of symptoms, suggest that this part of the palm is where phytoplasmas first move from leaves after foliar feeding by vectors and in which they probably multiply and distribute to other palm parts, including roots. Embryos from infected palms were analysed by nested‐PCR and 28% of 394 embryos were positive. Phytoplasma DNA was detected in embryos from fruit on any of the fruiting bunches regardless the age, but no pattern of quantitative distribution throughout the bunch developmental stages was observed. Germination of seeds from LY‐positive symptomatic palms was 58% and from LY‐negative symptomless palms were 71%. No phytoplasma was detected in seedlings tested from both symptomatic and non‐symptomatic palms. Seedlings tested after 2 years did not develop LY symptoms or eventually died.     

Growth patterns in the stem of the palm Archontophoenix cunninghamiana

Studies on the growth of Archontophoenix cunninghamiana in two separate populations show that this species establishes only a narrow apical meristem complex before rising above ground This complex continues to expand in girth as it rises, until at a height of about 12 m its girth becomes fixed. The lower regions of the trunk expand, more than equalling the expansion at the apex, producing at least a 20-fold increase in volume. Anatomical observations show that this expansion, termed sustained primary growth , is due to cell enlargement and cell division in the ground tissue, and is relatively uniform along the radius of the stem. When the apex reaches a height of 12 m, sustained primary growth ceases abruptly, fixing the taper of the trunk. Subsequent growth produces a cylindrical top on this tapered base.
These results are considered in the light of a review of earlier work on palm growth. The determinate nature of this type of arborescence is discussed, and brief reference is made to its evolutionary status.     

Organization of the vascular system in the stems of Diplazium and Blechnum (Filicales)

Detailed analysis of the three-dimensional vascular organization in species of Diplazium and Blechnum indicates that these ferns possess reticulate (dictyostelic) vascular systems that closely reflect the helical phyllotaxis of the shoot. In each species, the vascular pattern shows a specific relationship to the phyllotaxis, so that the phyllotactic fraction can be determined by examination of the number of cauline vascular bundles (meristeles) in cross section of the stem. The number of meristeles in a cross section equals the denominator of the phyllotactic fraction, i.e., the number of foliar orthostichies on the stem. The same numerical relationship also exists in the eusteles of seed plants between the number of axial (sympodial) stem bundles and the phyllotaxis. There is a further parallel between the three-dimensional reticulate pattern of fern dictyosteles and the reticulate patterns that characterize some herbaceous dicotyledons. However, the hypothesized separate origins of seed plant eusteles and fern dictyosteles from protostelic precursors preclude any direct homologies between these similar patterns. The parallel evolution of presumably more physiologically efficient reticulate systems in herbaceous seed plants and in ferns that have only a primary plant body is noteworthy. The similar relationships between the primary stem vascular patterns and phyllotaxy in both ferns and seed plants further emphasize the likely similarity of the morphogenetic events that occur at the shoot apex in these taxonomically disparate groups.     

The impact of utilization of palm products on the population structure of the vegetable ivory palm (Hyphaene petersiana,arecaceae) in North-Central Namibia

Indigenous trees fulfil many subsistence and economic needs in north-central Namibia.Hyphaene petersiana provides a range of products which contribute to most aspects of people’s livelihoods. Of particular importance is its income-generating capacity through the use of palm leaves for basket production and the sale of liquor distilled from the fruits. This study investigates the population structure ofHyphaene petersiana in two areas of different human and livestock densities. Data were recorded for height class distribution, basal diameter of mature, stemmed individuals and sex ratios. These parameters of population structure indicate a reduction in the recruitment of mature palms and an increase in single-stemmed, vegetatively reproduced palm suckers of the smallest size class (0.5 m). This trend is more pronounced in the site with greater human and livestock population densities. It appears to be related to high recorded levels of browsing by livestock of juvenile, unstemmed palms, despite the unpalatability of palm leaves. This acts to prevent recruitment into larger size classes and increase the compensatory growth of palm suckers, the latter being enhanced due to reduced competition through the prior removal by grazing animals of grasses and other herbaceous species. Accompanying this heavy pressure on juvenile palms are destructive uses of mature, stemmed palms, including their felling for construction purposes and tapping for palm wine. With regional human population increase, exacerbated by a recent trend to privatise land and raise pressure on remaining communal resources, it is possible that these destructive uses of mature palms will increase to unsustainable levels. Concern is thus expressed in this study regarding the long-term viability ofHyphaene petersiana populatiops in this area.     

Phyllotaxis in the Oil Palm: Application in the Evaluation of pisiferas

Yield differences between and within four segregating familiesof Oil Palm are shown to be highly correlated with differencesin the phyllotaxis index, E.P.I., of the female inflorescences.This association should facilitate prediction of the yield potentialof an effectively sterile segregant palm type known as pisifera,which is used as a male parent in hybrid seed production.     

Hormonal effects on phyllotaxis ofEuphorbia lathyris L.

Benzyladenine and gibberellic acid sprays (100 mg/l) onEuphorbia lathyris L. plants were absolutely successful (100% of the plants) in inducing a change in its normal decussate phyllotaxis. Benzyladenine produced a change to tetracussate, tricussate and bijugate, and gibberellic acid to spiral phyllotaxis. Benzyladenine and gibberellic acid treatments resulted in a significant increase in apex diameter (72.8% and 19.1% respectively). CCC, Ancymidol, Alar and Glyphosine did not alter decussate phyllotaxis.     

Characterization of Growth-Retardant Effects on Vegetative Growth of Date Palm Seedlings

The leaves, inflorescences, and fruit bunches of date palm trees develop at the crown, which is the apex of a single trunk that can be up to 25 m tall. Because palm trees are monocots, the leaves and inflorescences are generated deep within the trunk from a single meristem—the palm heart—and are inaccessible for direct bioregulator treatments. We followed the effects of two growth regulators, paclobutrazol and uniconazole-P, on the vegetative growth of young date palm seedlings. Both growth regulators constrained the elongation rate of the leaves and reduced the leaf length. Microscopic analysis of dissected palm hearts revealed that the growth retardants did not affect the generation of new leaves. However, the treatments did change the dimensions of the developing embryonic leaves as well as the shape of the apical meristem. The results suggest the possible future use of growth retardants to reduce trunk height, enabling cheaper, safer, and more efficient management of date palm trees.