A SURVEY OF THE VEGETATIVE SHOOT APICES IN THE FAMILY MALVACEAE

The present study compares the structure of the vegetative shoot apex in 40 species of the Malvaceae. There is a wide range of size, shape, and zonation within the apices of the family. Although many of the apices are domed, some are flat-topped and do not extend above the axil of the youngest leaf primordium. Also, most of the species investigated are recorded as having a more or less marked cytohistological zonation superimposed on the tunica-corpus configuration. The tunica is single-layered in a majority of species, but stratification of the upper corpus is common. In an effort to give a more accurate concept of apical structure and activity, the apex is described as the metrameristem and its derivatives: the flanking meristem, and the pith rib meristem or pith mother cells. The metrameristem, consisting of the tunica initials and the co pus initials, is the focal point of the study of the zoned apices. Data are presented for the measurements of the metrameristem, measurements of the apical dome, type of flanking meristem, origin of the pith, and growth habit of the plant. There appears to be a correlation between growth habit and the distinctness with which the metrameristem is marked off from the surrounding tissue. Most of the herbaceous species have an indistinctly marked metrameristem, whereas the shrubby trees and trees have a distinctly marked metrameristem. Zonation in shrubs and suffrutescent plants may be of either type.     

Variation in nicotine content of tobacco callus cultures

Plants ofNicotiana tabacum L. cv. Burley 21 which showed no difference in nicotine content were used to establish callus cultures. Cultures were initiated from different plants and from different leaves within each plant. The nicotine content of the calli was determined, and the results subjected to an analysis of variance. Differences between plants and differences within plants significantly affected the nicotine content of the cultures. The differences between plants were transmitted sexually and asexually, providing evidence that they are genetically determined. No such differences in nicotine content were found between the plants from which the cultures were established, indicating that nicotine production in vitro involves additional genes to those which are needed for nicotine production in the plant. The differences within plants were further investigated by establishing callus cultures from pith explants taken from different parts of the stem. Explants from apical pith tissue gave calli having far more nicotine and more roots than cultures derived from basal pith explants. This results may reflect the proximity of the apical pith explants to the site of auxin synthesis in the stem apex. Callus cultures derived from pith explants showed greater growth and nicotine production than those derived from leaf explants when the calli were induced on Murashige-Skoog medium containing -naphthalene acetic acid. This result is in conflict with the widely held belief that explants from different parts of the plant give cultures with similar yields of species-specific compounds.Abbreviations HN High nicotine - LN low nicotine     

No evidence of a generalized potential ‘cost’ of apical dominance for species that have strong apical dominance

尚无证据表明顶端优势强的物种存在广义顶端优势潜在“成本”     

Growth from two transient apical initials in the meristem of Selaginella kraussiana

A major transition in land plant evolution was from growth in water to growth on land. This transition necessitated major morphological innovations that were accompanied by the development of three-dimensional apical growth. In extant land plants, shoot growth occurs from groups of cells at the apex known as meristems. In different land plant lineages, meristems function in different ways to produce distinct plant morphologies, yet our understanding of the developmental basis of meristem function is limited to the most recently diverged angiosperms. To redress this balance, we have examined meristem function in the lycophyte Selaginella kraussiana. Using a clonal analysis, we show that S. kraussiana shoots are derived from the activity of two short-lived apical initials that facilitate the formation of four axes of symmetry in the shoot. Leaves are initiated from just two epidermal cells, and the mediolateral leaf axis is the first to be established. This pattern of development differs from that seen in flowering plants. These differences are discussed in the context of the development and evolution of diverse land plant forms.     

Cell-lineage patterns in the shoot apical meristem of the germinating maize embryo

A fate map for the shoot apical meristem of Zea mays L. at the time of germination was constructed by examining somatic sectors (clones) induced by -rays. The shoot apical meristem produced stem, leaves, and reproductive structures above leaf 6 after germination and the analysis here concerns their formation. On 160 adult plants which had produced 17 or 18 leaves, 277 anthocyanin-deficient sectors were scored for size and position. Sectors found on the ear shoot or in the tassel most often extended into the vegetative part of the plant. Sectors ranged from one to six internodes in length and some sectors of more than one internode were observed at all positions on the plant. Single-internode sectors predominated in the basal internodes (7,8,9) while longer sectors were common in the middle and upper internodes. The apparent number of cells which gave rise to a particular internode was variable and sectors were not restricted to the lineage unit: a leaf, the internode below it, and the axillary bud and prophyll at the base of the internode. These observations established two major features of meristem activity: 1) at the time of germination the developmental fate of any cell or group of cells was not fixed, and 2) at the time of germination cells at the same location in a meristem could produce greatly different amounts of tissue in the adult plant. Consequently, the developmental fate of specific cells in the germinating meristem could only be assigned in a general way.Abbreviations ACN apparent cell number - LI, LII, LI-LII sectors restricted to the epidermis, the subepidermis, or encompassing epidermis and subepidermis - PCN progenitor cell     

Effects of turgor potentials of epidermal cells neighbouring guard cells on stomatal opening in detached leaf epidermis and intact leaflets of Vicia faba L. (faba bean)

Leaflets of Vicia faba L. (faba bean) were used to determine whether the mechanical forces resulting from the turgor potentials (Φ_p) of the larger epidermal cells neighbouring guard cells play a significant role in regulating stomatal aperture. When Φ_p, of epidermis and Φ_p of bulk leaflet tissue were compared at midday, Φ_p of epidermis were only 15–25% those of bulk leaflet tissue at all but the most negative leaflet water potentials (Φ). When plants were bagged to increase Φ by reducing vapour pressure differences between leaflets and air, Φ_p of bulk leaflet tissue increased to predawn values, but Φ_p, of epidermis increased to only = 20% of predawn values and stomata opened to their widest apertures. Stomatal apertures were positively correlated with Φ_p of bulk leaflet tissue but they were not correlated with Φ_p of epidermis. Reductions in epidermal Φ_p, began predawn, before stomata were open, and reached minimum values at midday, when stomata were open. We conclude that, in Vicia faba, (1) reduction of Φ_p of epidermal cells begins predawn, reducing the counterforce to stomatal opening that would exist if full epidermal turgor were maintained throughout the day, and (2) changes in Φ_p, of leaf epidermal cells do not play a significant role in regulating stomatal aperture.     

AN INVESTIGATION OF THE PHYLOGENETIC AND ONTOGENETIC VARIABILITY OF SHOOT APICAL MERISTEMS IN THE CACTACEAE

The sizes, shapes and zonations of the shoot apical meristems of 22 species of cacti were examined. This family was chosen because of its great diversity of habits; the more primitive members are nonsucculent. leafy trees and more advanced members are highly specialized “leaf-less” stem-succulents. By combining these measurements with those already in the literature, a sample of almost 70 species was obtained. Apical meristems range in size from only 80 μm in diam in some species to as much as 1.500 μm in diam in others. The shape ranges from being flat to almost hemispherical. Despite the great range in size and shape of the apical meristems, or the range in the morphologies of the leaves and stems which are produced by the meristems. all apices had the usual zonation: tunica, central mother cells, peripheral zone, and pith-rib meristem. The sizes of each of the zones. expressed either as the number of cells per zone or expressed as a percentage of the whole apex. were highly variable. The variation in apical dimensions and zone sizes occurred both phylogenetically and ontogenetically. and this has been interpreted to indicate that the morphogenetic mechanisms which control apical size and zonation are easily modified, both during the development of individual plants and during the evolution of new species.     

Herbivory could unlock mutations sequestered in stratified shoot apices of genetic mosaics

Many higher plants have shoot apical meristems that possess discrete cell layers, only one of which normally gives rise to gametes following the transition from vegetative meristem to floral meristem. Consequently, when mutations occur in the meristems of sexually reproducing plants, they may or may not have an evolutionary impact, depending on the apical layer in which they reside. In order to determine whether developmentally sequestered mutations could be released by herbivory (i.e., meristem destruction), a characterized genetic mosaic was subjected to simulated herbivory. Many plants develop two shoot meristems in the leaf axils of some nodes, here referred to as the primary and secondary axillary meristems. Destruction of the terminal and primary axillary meristems led to the outgrowth of secondary axillary meristems. Seed derived from secondary axillary meristems was not always descended from the second apical cell layer of the terminal shoot meristem as is expected for terminal and primary shoot meristems. Vegetative and reproductive analysis indicated that secondary meristems did not maintain the same order of cell layers present in the terminal shoot meristem. In secondary meristems reproductively sequestered cell layers possessing mutant cells can be repositioned into gamete-forming cell layers, thereby adding mutant genes into the gene pool. Herbivores feeding on shoot tips may influence plant evolution by causing the outgrowth of secondary axillary meristems.     

VARIATION IN APPLE CYTOCHIMERAS

Diploid-tetraploid cytochimeras of apple were investigated by cytological examination of buds from branches selected by the characteristics of the fruits, flowers or pollen. Eight types of cytochimeras were identified on the basis of relative size of cells, nuclei and metaphase mitoses in the apical meristem (protomeristem) and mitoses in meristematic primary tissues. There appeared to be five apical layers which contributed to the stem and four which contributed to the leaves and flower parts. Buds of various cytochimeral patterns (designated by a formula giving the ploidy of the apical layers) were found on some bearing trees propagated from known cytochimeral sources. The most frequently associated types were (a) 2–4–2–2–2 and 2–4–4–4–4 (and 2–4–2–4–4 in one cultivar), or (b) 2–2–2–4–4 and 2x. Some sports were uniformly 2–4–4–4–4. The stability of apple cytochimeras under normal conditions appeared greater in some eultivars than in others. Sprouts from severely pruned 2–2–4–4–4 trees were more variable than unpruned branches. Buds of shoots which grew from radiation-damaged buds were more variable than those from non-irradiated buds and included types not yet found by branch selection. Cytochimeral variation was interpreted to be due to layer replacement resulting from infrequent periclinal divisions in apical or axillary meristems, or from wounding of meristems by ionizing radiation.     

Cell division and morphological changes in the shoot apex of Arabidopsis thaliana during floral transition

Eight-week-old vegetative plants of Arabidopsis thaliana, ecotype Columbia, were induced to flower by a single long day (LD). In this experimental system, it is known that the last component of the floral stimulus moves from the leaves to the apex 24-36 h after the start of the LD, and the first floral meristem is initiated by the shoot apical meristem (SAM) at 44-56 h (Corbesier et al., 1996, The Plant Journal 9: 947-952). Here we show that the rate of cell division is increased at floral transition in all SAM parts but not in the sub-apical pith cells. Mitotic activity starts to increase 24 h after the start of the LD and is two- to three-fold higher at peak times than that in non-induced plants. This activation is followed by the start of SAM enlargement at 44 h, SAM doming at 48 h, and the elongation of apical internodes (bolting) at 52 h.     

Apical meristem organization and lack of establishment of the quiescent center in Cactaceae roots with determinate growth

Some species of Cactaceae from the Sonoran Desert are characterized by a determinate growth pattern of the primary root, which is important for rapid lateral-root formation and seedling establishment. An analysis of the determinate root growth can be helpful for understanding the mechanism of meristem maintenance in plants in general. Stenocereus gummosus (Engelm.) Gibson & Horak and Pachycereus pringlei (S. Watson) Britton & Rose are characterized by an open type of root apical meristem. Immunohistochemical analysis of 5-bromo-2-deoxyuridine incorporation into S. gummosus showed that the percentage of cells passing through the S-phase in a 24-h period is the same within the zone where a population of relatively slowly proliferating cells could be established and above this zone in the meristem. This indicated the absence of the quiescent center (QC) in S. gummosus. During the second and the third days of growth, in the distal meristem portion of P. pringlei roots, a compact group of cells that had a cell cycle longer than in the proximal meristem was found, indicating the presence of the QC. However, later in development, the QC could not be detected in this species. These data suggest that during post-germination the absence of the establishment of the QC within the apical meristem and limited proliferative activity of initial cells are the main components of a determinate developmental program and that establishment of the QC is required for maintenance of the meristem and indeterminate root growth in plants.Abbreviations QC quiescent center - RCP root cap-protoderm - BrdU 5-bromo-2-deoxyuridine - FITC fluorescein isothiocyanate - DAPI 4,6-diamidino-2-phenylindole     

The multifaceted roles of FLOWERING LOCUS T in plant development

One of the key developmental processes in flowering plants is the differentiation of the shoot apical meristem into a floral meristem. This transition is regulated through the integration of environmental and endogenous stimuli, involving a complex, hierarchical signalling network. In arabidopsis, the FLOWERING LOCUS T (FT) protein, a mobile signal recognized as a major component of florigen, has a central position in mediating the onset of flowering. FT-like genes seem to be involved in regulating the floral transition in all angiosperms examined to date. Evidence from molecular evolution studies suggests that the emergence of FT-like genes coincided with the evolution of the flowering plants. Hence, the role of FT in floral promotion is conserved, but appears to be restricted to the angiosperms. Besides flowering, FT-like proteins have also been identified as major regulatory factors in a wide range of developmental processes including fruit set, vegetative growth, stomatal control and tuberization. These multifaceted roles of FT-like proteins have resulted from extensive gene duplication events, which occurred independently in nearly all modern angiosperm lineages, followed by sub- or neo-functionalization. This review assesses the plethora of roles that FT-like genes have acquired during evolution and their implications in plant diversity, adaptation and domestication.     

Understanding plant reproductive diversity

Flowering plants display spectacular floral diversity and a bewildering array of reproductive adaptations that promote mating, particularly outbreeding. A striking feature of this diversity is that related species often differ in pollination and mating systems, and intraspecific variation in sexual traits is not unusual, especially among herbaceous plants. This variation provides opportunities for evolutionary biologists to link micro-evolutionary processes to the macro-evolutionary patterns that are evident within lineages. Here, I provide some personal reflections on recent progress in our understanding of the ecology and evolution of plant reproductive diversity. I begin with a brief historical sketch of the major developments in this field and then focus on three of the most significant evolutionary transitions in the reproductive biology of flowering plants: the pathway from outcrossing to predominant self-fertilization, the origin of separate sexes (females and males) from hermaphroditism and the shift from animal pollination to wind pollination. For each evolutionary transition, I consider what we have discovered and some of the problems that still remain unsolved. I conclude by discussing how new approaches might influence future research in plant reproductive biology.     

Shoot meristem formation and maintenance

The shoot apical meristem of higher plants is a self-maintaining stem cell system which gives rise to the entire above-ground part of a plant. In the past year, genetic and molecular studies have provided increasing insight into the processes of shoot meristem formation and maintenance, as well as into the relation between the apical meristem and its products.     

<Emphasis Type="Italic">ZWILLE</Emphasis> buffers meristem stability in<Emphasis Type="Italic"> Arabidopsis thaliana</Emphasis>

The shoot apical meristem of higher plants consists of a population of stem cells at the tip of the plant body that continuously gives rise to organs such as leaves and flowers. Cells that leave the meristem differentiate and must be replaced to maintain the integrity of the meristem. The balance between differentiation and maintenance is governed both by the environment and the developmental status of the plant. In order to respond to these different stimuli, the meristem has to be plastic thus ensuring the stereotypic shape of the plant body. Meristem plasticity requires the ZWILLE (ZLL) gene. In zll mutant embryos, the apical cells are misspecified causing a variability of the meristems size and function. Using specific antibodies against ZLL, we show that the zll phenotype is due to the complete absence of the ZLL protein. In immunohistochemical experiments we confirm the observation that ZLL is solely localized in vascular tissue. For a better understanding of the role of ZLL in meristem stability, we analysed the genetic interactions of ZLL with WUSCHEL (WUS) and the CLAVATA1, 2 and 3 (CLV) genes that are involved in size regulation of the meristem. In a zll loss-of-function background wus has a negative effect whereas clv mutations have a positive effect on meristem size. We propose that ZLL buffers meristem stability non-cell-autonomously by ensuring the critical number of apical cells required for proper meristem function.Edited by G. JürgensAn erratum to this article can be found at     

Changes of axillary meristem localization in relation to flowering ofRaphanus sativus L. after GA treatment

Two phases of radish ontogenesis (I-when the plant had produced 3 –5 nodes and II-when the plant had produced 8 –10 nodes) were established on the basis of axillary, meristem localization. Flowering of the plants in response to GA treatment depends on the phases in which they were treated and on growth correlations in the apical meristem. The results obtained suggest that the reaction ofRaphanus sativus (LDP) to GA treatment is parallel to that ofChenopodium rubrum (SDP), and that the response of radish plants also depends on changes in growth correlations in the shoot apical meristem at the time of treatment.     

The nutritional status of the apical meristem of Lactuca sativa as affected by NaCl salinization: An electron-probe microanalytic study

A volume of tissue of lettuce (Lactuca sativa L.) plants extending 2 mm basipetally from the apical meristem and including leaf primordia and young expanding leaves was surveyed using electron-probe microanalysis (EPMA) on both frozen-hydrated and freeze-dried samples. This analysis was carried out either 2 or 5 d following NaCl salinization of the medium from the 10 mol · m^–-3 control level up to 80 mol · m^–-3. The objective was the investigation of possible changes in the nutritional status of the apical meristem that might account for some aspects of salt-induced growth inhibition. Sodium and chloride increased significantly in tissues basal to the apical meristem, while both phosphorus and potassium decreased in the same region. These changes were evident in specimens collected just 2 d after the commencement of salinization (20 h after completion of the salinization) and were not exacerbated by an additional 3 d of treatment; they were present in tissue as close as 100 m to the meristem and extending down to 500 m. The apical 10–50 m were relatively protected from both the increase in sodium and chloride and the decrease in phosphorus and potassium that occurred in more basal regions. Young leaves (up to 1.5 mm in length) appear to control their own mineral nutrient levels when challenged by salinization of the medium, presumably because of altered growth. A decrease in the concentration of total Ca as a result of salinization was significant in cells 500 m basal to the meristem, but was evident as a tendency in the data even within the first 50 m. Using an improved automatic method for the analysis of calcium by EPMA, it was found that total Ca was reduced by salinization, especially in basal regions (500 m below the apex) and also in young leaves (1–1.5 mm in length). We suggest that the nutrition of the shoot apical meristem may be disturbed soon after salinization and that the shoot meristem might be the source of a signal to expanding leaves, as well as exerting its own direct influence over leaf emergence.Abbreviation EPMA electron-probe microanalysis This work was supported by U.S. Department of Agriculture grant 87-CRCR-1-2462.     

HIV-1 proteins preferentially activate anti-inflammatory M2-type macrophages

HIV-1 proteins, including Tat, gp120, and Nef, activate macrophages (MΦ), which is consistent with the fact that HIV-1 infection is characterized by sustained immune activation. Meanwhile, MΦ are functionally classified into two types: proinflammatory M1-MΦ and anti-inflammatory M2-MΦ. We show that HIV-1 proteins, particularly Nef, preferentially activate M2-MΦ. Extracellular Tat, gp120, and Nef activated MAPK and NF-κB pathways in human peripheral blood monocyte-derived MΦ. However, the activation was marked in M-CSF-derived M2-MΦ but not GM-CSF-derived M1-MΦ. Nef was the most potent activator, and its signaling activation was comparable to that by TNF-α. Indeed, Nef was internalized more rapidly by M2-MΦ than by M1-MΦ. The myristoylation and proline-rich motif of Nef were responsible for the observed signaling activation. Consistent with the activation of MAPK/NF-κB pathways, Nef stimulated the production of a number of proinflammatory cytokines/chemokines by M2-MΦ. However, Nef reduced the expression of CD163 and phagocytosis, the characteristic markers of M2-MΦ, indicating that Nef drives an M2-like to M1-like phenotypic shift. Because the differentiation of most tissue MΦ depends on M-CSF and its receptor, which is the essential axis for the anti-inflammatory M2-MΦ phenotype, the current study reveals an efficient mechanism by which HIV-1 proteins, such as Nef, induce the proinflammatory MΦ.     

Effects of apical damage on plant growth and male and female reproductive investments in <Emphasis Type="Italic">Ambrosia artemisiifolia</Emphasis>, a wind-pollinated plant

In wind-pollinated plants, apical damage may decrease male fitness by reducing height-dependent pollen dispersal distance, but may not affect female fitness because plant height is not always correlated with female fitness. We hypothesized that Ambrosia artemisiifolia responds to apical damage by (1) restoring plant height through compensatory growth from lateral buds, and/or (2) increasing the sex allocation to female function to compensate for the loss of male fitness. We tested these hypotheses by comparing a group of experimental removal of the apical meristem with three control groups and by field surveys on apically damaged plants. Experimental apical damage suppressed main stem growth, but promoted vertical secondary growth from lateral buds. These responses resulted in compensation of stem height in the apically damaged plants to the same height as one of three control groups. The numbers of male and female flowers and male racemes did not differ between damaged and undamaged plants, indicating that apically damaged plants did not change their sex allocation. Therefore, our results support our first hypothesis. The results of a field survey of naturalized populations also supported the first hypothesis in that plant height and the number of male racemes did not change in plants with apical damage. Consequently, our results suggest that A. artemisiifolia has a high ability of fitness compensation after apical damage by restoring height and male function. This ability may contribute to its invasiveness in disturbed habitats.