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Phytochromes are dimeric proteins that function as red and far-red light sensors influencing nearly every phase of the plant life cycle. Of the three major phytochrome families found in flowering plants, PHYTOCHROME C (PHYC) is the least understood. In Arabidopsis and rice, PHYC is unstable and functionally inactive unless it heterodimerizes with another phytochrome. However, when expressed in an Arabidopsis phy- null mutant, wheat PHYC forms signaling active homodimers that translocate into the nucleus in red light to mediate photomorphogenic responses. Tetraploid wheat plants homozygous for loss-of-function mutations in all PHYC copies (phyC ᴬᴮ) flower on average 108 d later than wild-type plants under long days but only 19 d later under short days, indicating a strong interaction between PHYC and photoperiod. This interaction is further supported by the drastic down-regulation in the phyC ᴬᴮ mutant of the central photoperiod gene PHOTOPERIOD 1 (PPD1) and its downstream target FLOWERING LOCUS T1 , which are required for the promotion of flowering under long days. These results implicate light-dependent, PHYC-mediated activation of PPD1 expression in the acceleration of wheat flowering under inductive long days. Plants homozygous for the phyC ᴬᴮ mutations also show altered profiles of circadian clock and clock-output genes, which may also contribute to the observed differences in heading time. Our results highlight important differences in the photoperiod pathways of the temperate grasses with those of well-studied model plant species.

Selaginella species are characterized by regular anisotomous dichotomous divisions of the shoot apical meristem, giving rise to two new axes (branches) which differ in size. A vital process is the formation of vascular connections, which enables continuous communication and consequent functional and developmental integration of a plant during branching. Here, we present the sequence of developmental changes in the vascular system of Selaginella kraussiana related to dichotomous branching. Stem vasculature in Selaginella kraussiana consists of two meristeles which change in arrangement during shoot development. Using dye tracers, we documented developmental functional isolation of meristeles associated with the specific structure of the stelar system, which results in a spatiotemporal sectoriality of the shoot. We discuss sectoriality in terms of possible significance for shoot development.

The response of the stem of a resistant host (Impatiens baslamina) to infection by the parasitic flowering plant Cuscuta japonica was studied with light and electron microscopy. The intra- and interfascicular cambial cells in the host stem first reacted to the penetrating upper haustorium by dividing, and the differentiation of the host xylem (vascular) tissues proceeded toward interfascicular areas from vascular bundles. When the host vascular tissue was invaded by the endophyte (haustorial portion in the host stem), the host xylem was displaced, and host vessels became occluded with parenchyma cells, resulting in tyloses. As the parasitism progressed, areas of the host stem penetrated by the endophyte became swollen via secondary growth and cell division in the parenchymatous cortex, pith, and interfascicular areas. During this intrusion by the endophyte, darkly stained necrotic reactions were detected at the interface between the host tissue and the invading endophyte. The results suggested that in the host tissues penetrated by the parasite, the formation of secondary tissue and swellings caused by active cell division of ground tissue and host vessel occlusion by tyloses constitute the host structural defense against the parasite.

Enhancing nutrient uptake and the subsequent elemental transport from the sites of application to sites of utilization is of great importance to the science and practical field application of foliar fertilizers. The aim of this study was to investigate the mobility of various foliar applied zinc (Zn) formulations in sunflower (Helianthus annuus L.) and to evaluate the effects of the addition of an organic biostimulant on phloem loading and elemental mobility. This was achieved by application of foliar formulations to the blade of sunflower (H. annuus L.) and high-resolution elemental imaging with micro X-ray fluorescence (μ-XRF) to visualize Zn within the vascular system of the leaf petiole. Although no significant increase of total Zn in petioles was determined by inductively-coupled plasma mass-spectrometer, μ-XRF elemental imaging showed a clear enrichment of Zn in the vascular tissues within the sunflower petioles treated with foliar fertilizers containing Zn. The concentration of Zn in the vascular of sunflower petioles was increased when Zn was applied with other microelements with EDTA (commercial product Kick-Off) as compared with an equimolar concentration of ZnSO4 alone. The addition of macronutrients N, P, K (commercial product CleanStart) to the Kick-Off Zn fertilizer, further increased vascular system Zn concentrations while the addition of the microbially derived organic biostimulant \"GroZyme\" resulted in a remarkable enhancement of Zn concentrations in the petiole vascular system. The study provides direct visualized evidence for phloem transport of foliar applied Zn out of sites of application in plants by using μ-XRF technique, and suggests that the formulation of the foliar applied Zn and the addition of the organic biostimulant GroZyme increases the mobility of Zn following its absorption by the leaf of sunflower.

Structural changes in apical meristems accompany morphological specialization and adaptation of vascular-plant sporophytes to changing environments. Analysis of cell clones at the shoot apical meristem surface of a representative of the basal vascular plants made it possible to trace the dynamics of initial cells, which are recorded in cell wall arrangements. The cellular pattern at the summit of the Huperzia lucidula (Michx.) Trevisan shoot apical meristem is usually composed of four initial cells, forming a tetrad at the apex center, and their derivative clones. Our data show that the apical initial cells present at a given time in a tetrad often are transient and replaced by new initials in the ontogeny of a given Huperzia apex. Specifically, the process of dichotomous branching leads to extensive cell division, resulting in a pool of potential initial cells from which the new initial centers of the twin axes are selected. The ability to specify newly arisen cells as initials during shoot dichotomy indicates that a mechanism for selection and replacement of initial cells may exist in lycopods. Such a cellular mechanism, which could play an important role in protecting the genetic identity and integrity of meristematic cells, would be expected to emerge early in the phylogeny of vascular plants.

Ginkgo biloba, the only living representative in an otherwise extinct clade, is of pivotal importance to understanding seed plant phylogeny. Although G. biloba and its fossil relatives have been studied for over two centuries, there are both gaps and contradictions in the information available. We present data documenting the distributions of strobili and consider what an understanding of the disposition of strobili along short-shoots in Ginkgo adds to knowledge of the evolution of reproductive structures in seed plants in general. The megasporangiate strobili are found at and around the boundary between bracts and foliage leaves, while the expanse of microsporangiate strobili centers on the fifth bract back from that boundary. Quantitative analysis of the locations of the strobili along the short-shoot finds that increases in numbers of strobili are the result of recruitment of adjacent axils into morphogenetic activity. Gaps in the series of strobili are exceedingly rare. Further, while increased numbers of megasporangiate strobili arise from the symmetrical addition of axils into the fertile zone, increased numbers of microsporangiate strobili arise from a distinctly asymmetrical, basipetally biased, addition of axillary positions. This accurate morphological framework should orient molecular genetic studies that probe gymnosperm development itself or that consider gymnosperms as the proximate sources of gene expression redeployed in the origin of the angiosperm flower.

• Premise: Ginkgo, centrally placed in seed plant phylogeny, is considered important in many phylogenetic and evolutionary studies. Shoot dimorphism of Ginkgo has been long noted, but no work has yet been done to evaluate the relationships between overall branch architecture and wood ring characters, shoot growth, and environmental conditions.• Methods: Branches, sampled from similar canopy heights, were mapped with the age of each long shoot segment determined by counting annual leaf-scar series on its short shoots. Transverse sections were made for each long shoot segment and an adjacent short shoot; wood ring thickness, number of rings, and number of tracheids/ring were determined. Using branch maps, we identified wood rings for each long shoot segment to year and developmental context of each year (distal short shoot growth only vs. at least one distal long shoot). Climate data were also analyzed in conjunction with developmental context.• Key results: Significantly thicker wood rings occur in years with distal long shoot development. The likelihood that a branch produced long shoots in a given year was lower with higher maximum annual temperature. Annual maximum temperature was negatively correlated with ring thickness in microsporangiate trees only. Annual minimum temperatures were correlated differently with ring thickness of megasporangiate and microsporangiate trees, depending on the developmental context. There were no significant effects associated with precipitation.• Conclusions: Overall, developmental context alone predicts wood ring thickness about as well as models that include temperature. This suggests that although climatic factors may be strongly correlated with wood ring data among many gymnosperm taxa, at least for Ginkgo, correlations with climate data are primarily due to changes in proportions of shoot developmental types (LS vs. SS) across branches.

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.

Structural changes in apical meristems accompany morphological specialization and adaptation of vascular-plant sporophytes to changing environments. Analysis of cell clones at the shoot apical meristem surface of a representative of the basal vascular plants made it possible to trace the dynamics of initial cells, which are recorded in cell wall arrangements. The cellular pattern at the summit of theHuperzia lucidula(Michx.) Trevisan shoot apical meristem is usually composed of four initial cells, forming a tetrad at the apex center, and their derivative clones. Our data show that the apical initial cells present at a given time in a tetrad often are transient and replaced by new initials in the ontogeny of a givenHuperziaapex. Specifically, the process of dichotomous branching leads to extensive cell division, resulting in a pool of potential initial cells from which the new initial centers of the twin axes are selected. The ability to specify newly arisen cells as initials during shoot dichotomy indicates that a mechanism for selection and replacement of initial cells may exist in lycopods. Such a cellular mechanism, which could play an important role in protecting the genetic identity and integrity of meristematic cells, would be expected to emerge early in the phylogeny of vascular plants.

Rhizophores arise from angle meristems located at shoot branch points in many species of Selaginella and produce roots at their distal ends. We reinvestigated the origin and development of rhizophores and roots in Selaginella martensii using a nondestructive sequential replica technique for scanning electron microscopy and anatomical studies of sectioned material. We followed development in excised branch systems containing undetermined ventral angle meristems Analysis of patterns of division and expansion of cells on the surface of apical meristems and in transverse and longitudinal sections showed that the outgrowth from a ventral angle meristem had three distinct developmental phases: rhizophore, aerial root, and subterranean root, each with a distinctive apical meristem structure and activity. Rhizophore formation involved division of all cells in the angle meristem and establishment of multiple surface initial cells from which rhizophore growth ensued. The aerial root phase began with formation of two internal root meristems, each with a single prominent apical cell, replacing the multiple surface initials. From these meristems, dichotomously branched aerial roots formed, which continued to grow and branch internally. Eventually, subterranean roots formed, bearing root hairs and root caps. Formation of subterranean roots was induced at any time after aerial roots reached 0.5-1 cm in length when the tip of the organ touched a substrate. Our data provided support for the original concept of the rhizophore as a unique root-bearing organ in S. martensii, as well as for a developmental succession of apical meristem structure and activity, specific forms of which characterized the rhizophore and true root stages of angle meristem outgrowth.