Verbena officinalis Verbenaceae (Lamiales): a new plant model system for phyllotaxis research.

Phyllotactic diversity and developmental transitions between phyllotactic patterns are not fully understood. The plants studied so far, such as Magnolia, Torreya or Abies, are not suitable for experimental work, and the most popular model plant, Arabidopsis thaliana, does not show sufficient phyllotactic variability. It has been found that in common verbena (Verbena officinalis L.), a perennial, cosmopolitan plant, phyllotaxis differs not only between growth phases in primary transitions but also along the indeterminate inflorescence axis in a series of multiple secondary transitions. The latter are no longer associated with the change in lateral organ identity, and the sequence of phyllotactic patterns is puzzling from a theoretical point of view. Data from the experiments in silico, confronted with empirical observations, suggest that secondary transitions might be triggered by the cumulative effect of fluctuations in the continuously decreasing bract primordia size. The most important finding is that the changes in the primary vascular system, associated with phyllotactic transitions, precede those taking place at the apical meristem. This raises the question of the role of the vascular system in determining primordia initiation sites, and possibly challenges the autonomy of the apex. The results of this study highlight the complex relationships between various systems that have to coordinate their growth and differentiation in the developing plant shoot. Common verbena emerges from this research as a plant that may become a new model suitable for further studies on the causes of phyllotactic transitions.

Chiral events in developing gametophores of Physcomitrella patens and other moss species are driven by an unknown, universal direction-sensing mechanism.

PREMISE OF THE STUDY: We used the model species Physcomitrella patens to examine chirality in moss gametophores. Chirality is manifested in the direction of consecutive apical cell divisions, cell plate configurations, and deviations of leaf connecting lines from the vertical course. However, the frequencies of chiral configurations of all these processes as well as their mutual dependence-especially in the case of gametophore branching-are not known. Other moss species were checked to determine the universality of our findings. METHODS: The gametophore structure of Physcomitrella patens grown in the laboratory under controlled conditions was investigated using light microscopy and compared with that of other moss species collected from their natural stands. KEY RESULTS: In all investigated moss species, the tetrahedral apical cell exhibits either clockwise or counterclockwise consecutive divisions, and selection of this directionality in the primary axis is random. It is, however, related to cell plate configuration. If the plate is skewed, leaf-producing segments arising from the apical cell cleavage exhibit circumferential rotation. Three parallel lines connecting the leaves deviate from a vertical course, but always in the same direction as that of leaf initiation; thus, the angular distance between consecutive leaves increases to >120°. Lateral branches are exclusively antidromous. CONCLUSIONS: Gametophore chiral configuration appears to be useful in resolving problems of moss modular growth and branching. Morphological and anatomical evidence strongly suggests that an unknown direction-sensing mechanism controls the development of moss axial organs. We propose that leaves are responsible for a horizontal gradient of sugar signals that develops along the gametophore circumference, thus influencing branching-unit chirality.

Mechanochemical Polarization of Contiguous Cell Walls Shapes Plant Pavement Cells.

The epidermis of aerial plant organs is thought to be limiting for growth, because it acts as a continuous load-bearing layer, resisting tension. Leaf epidermis contains jigsaw puzzle piece-shaped pavement cells whose shape has been proposed to be a result of subcellular variations in expansion rate that induce local buckling events. Paradoxically, such local compressive buckling should not occur given the tensile stresses across the epidermis. Using computational modeling, we show that the simplest scenario to explain pavement cell shapes within an epidermis under tension must involve mechanical wall heterogeneities across and along the anticlinal pavement cell walls between adjacent cells. Combining genetics, atomic force microscopy, and immunolabeling, we demonstrate that contiguous cell walls indeed exhibit hybrid mechanochemical properties. Such biochemical wall heterogeneities precede wall bending. Altogether, this provides a possible mechanism for the generation of complex plant cell shapes.

Spatial pattern of long-distance symplasmic transport and communication in trees.

Symplasmic short- and long-distance communication may be regulated at different levels of plant body organization. It depends on cell-to-cell transport modulated by plasmodesmata conductivity and frequency but above all on morphogenetic fields that integrate a plant at the supracellular level. Their control of physiological and developmental processes is especially important in trees, where the continuum consists of 3-dimensional systems of: 1) stem cells in cambium, and 2) living parenchyma cells in the secondary conductive tissues. We found that long-distance symplasmic transport in trees is spatially regulated. Uneven distribution of fluorescent tracer in cambial cells along the branches examined illustrates an unknown intrinsic phenomenon that can possibly be important for plant organism integration. Here we illustrate the spatial dynamics of symplasmic transport in cambium, test and exclude the role of callose in its regulation, and discuss the mechanism that could possibly be responsible for the maintenance of this spatial pattern.

Symplasmic, long-distance transport in xylem and cambial regions in branches of Acer pseudoplatanus (Aceraceae) and Populus tremula x P. tremuloides (Salicaceae).

PREMISE OF THE STUDY: The picture of how long-distance transport proceeds in trees is still far from being complete. Beside the apoplasmic pathway, transport undoubtedly also takes place within the system of living cells in the secondary xylem and cambial region. Because detailed, thorough studies of the symplasmic routes in woody branches, using direct localization with fluorescent tracers, had not been done, here we focused on the main routes of long-distance symplasmic transport in xylem and cambial tissues and analyzed in detail tracer distribution in the rays on the extended cambial surface in branches of Acer pseudoplatanus and Populus tremula ×P. tremuloides. METHODS: Fluorescent tracers were loaded into branches through the vascular system, then their distribution in xylem and cambial regions was analyzed. KEY RESULTS: Tracer signal was present in the symplast of axial and radial xylem parenchyma cells and in both types of cambial cells. The living cells of xylem parenchyma and of the cambium were symplasmically interconnected via xylem rays. Tracer distribution in rays was uneven on the extended cambial surface; cambial regions with intensively or sparsely dyed rays alternated along the vertical axis of analyzed branches. CONCLUSIONS: Symplasmic, long-distance transport is present between the living cells of xylem and the cambial region in woody branches. The uneven distribution of fluorescent tracers in cambial rays along the stems is surprising and suggests the presence of an intrinsic pattern caused by an unknown mechanism.

Virtual phyllotaxis and real plant model cases.

Phyllotactic pattern results from genetic control of lateral primordia size (physiological or physical) relative to the size of organogenic lateral surface of shoot apical meristem (SAM). In order to understand the diversity of patterns and ontogenetic transitions of phyllotaxis we have developed a geometric model allowing changes of the above proportion in a computer simulation of SAM's growth. The results of serial simulations confirmed that many phyllotactic patterns (including most esoteric ones) and ontogenetic transitions known from real plant model cases can be easily obtained in silico. Properties of virtual patterns often deviated from those of ideal mathematical lattices but closely resembled those of the natural ones. This proved the assumptions of the model, such as initiation in the first available space or ontogenetic changes in primordia size, to be quite realistic. Confrontation of simulation results with some sequences of real phyllotactic patterns (case study Verbena) questions the autonomy of SAM in its organogenic activity and suggests the involvement of unknown signal positioning primordia in a non-random manner in the first available space.

Influence of a weak DC electric field on root meristem architecture.

BACKGROUND AND AIMS: Electric fields are an important environmental factor that can influence the development of plants organs. Such a field can either inhibit or stimulate root growth, and may also affect the direction of growth. Many developmental processes directly or indirectly depend upon the activity of the root apical meristem (RAM). The aim of this work was to examine the effects of a weak electric field on the organization of the RAM. METHODS: Roots of Zea mays seedlings, grown in liquid medium, were exposed to DC electric fields of different strengths from 0.5 to 1.5 V cm(-1), with a frequency of 50 Hz, for 3 h. The roots were sampled for anatomical observation immediately after the treatment, and after 24 and 48 h of further undisturbed growth. KEY RESULTS: DC fields of 1 and 1.5 V cm(-1) resulted in noticeable changes in the cellular pattern of the RAM. The electric field activated the quiescent centre (QC): the cells of the QC penetrated the root cap junction, disturbing the organization of the closed meristem and changing it temporarily into the open type. CONCLUSIONS: Even a weak electric field disturbs the pattern of cell divisions in plant root meristem. This in turn changes the global organization of the RAM. A field of slightly higher strength also damages root cap initials, terminating their division.

Vascular architecture in shoots of early divergent vascular plants, Lycopodium clavatum and Lycopodium annotinum.

Lycopodium represents a phylogenetically distinct clade of basal vascular plants with anatomical characters that have no parallel in other lineages. Thus, knowledge of lycopod structure and development may reveal important information about the common ancestors of all vascular plants. Here we report the unique architecture of the conducting system in Lycopodium annotinum and Lycopodium clavatum. Based on multiple series of anatomical sections, we reconstructed spatial relationships between microphylls and the stelar system. Analysis revealed that protoxylem ribs (PXR) were vertical, regardless of type of phyllotaxis, and their numbers were variable. Microphyll traces (MTr) were randomly distributed between ribs, resulting in the absence of defined sympodia and varied lengths of MTr. Dichotomous branching contributed to additional features, for example occurrence of mesarch protoxylem, affecting stele structure and PXR numbers. Our data showed limited interrelationships between lycopod vasculature and microphyll phyllotaxis. This may suggest that both systems developed independently, then evolved together to form the integrated supply system. Thus vasculature in extant lycophytes may be less functionally efficient than in seed plants, where consistent leaf-trace lengths guarantee predictable energy utilization during ontogeny. Differences may result from the phylogenetically different origin of microphylls, and the level of vascular complexity.