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Although the evolution of spores was critical to the diversification of plants on land, sporogenesis is incompletely characterized for model plants such as Physcomitrium patens . In this study, the complete process of P. patens sporogenesis is detailed from capsule expansion to mature spore formation, with emphasis on the construction of the complex spore wall and proximal aperture. Both diploid (sporophytic) and haploid (spores) cells contribute to the development and maturation of spores. During capsule expansion, the diploid cells of the capsule, including spore mother cells (SMCs), inner capsule wall layer (spore sac), and columella, contribute a locular fibrillar matrix that contains the machinery and nutrients for spore ontogeny. Nascent spores are enclosed in a second matrix that is surrounded by a thin SMC wall and suspended in the locular material. As they expand and separate, a band of exine is produced external to a thin foundation layer of tripartite lamellae. Dense globules assemble evenly throughout the locule, and these are incorporated progressively onto the spore surface to form the perine external to the exine. On the distal spore surface, the intine forms internally, while the spiny perine ornamentation is assembled. The exine is at least partially extrasporal in origin, while the perine is derived exclusively from outside the spore. Across the proximal surface of the polar spores, an aperture begins formation at the onset of spore development and consists of an expanded intine, an annulus, and a central pad with radiating fibers. This complex aperture is elastic and enables the proximal spore surface to cycle between being compressed (concave) and expanded (rounded). In addition to providing a site for water intake and germination, the elastic aperture is likely involved in desiccation tolerance. Based on the current phylogenies, the ancestral plant spore contained an aperture, exine, intine, and perine. The reductive evolution of liverwort and hornwort spores entailed the loss of perine in both groups and the aperture in liverworts. This research serves as the foundation for comparisons with other plant groups and for future studies of the developmental genetics and evolution of spores across plants.more » « less
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null (Ed.)A robust spore wall was a key requirement for terrestrialization by early plants. Sporopollenin in spore and pollen grain walls is thought to be polymerized and cross-linked to other macromolecular components, partly through oxidative processes involving H 2 O 2 . Therefore, we investigated effects of scavengers of reactive oxygen species (ROS) on the formation of spore walls in the moss Physcomitrella patens (Hedw.) Bruch, Schimp & W. Gümbel. Exposure of sporophytes, containing spores in the process of forming walls, to ascorbate, dimethylthiourea, or 4-hydroxy-TEMPO prevented normal wall development in a dose, chemical, and stage-dependent manner. Mature spores, exposed while developing to a ROS scavenger, burst when mounted in water on a flat slide under a coverslip (a phenomenon we named “augmented osmolysis” because they did not burst in phosphate-buffered saline or in water on a depression slide). Additionally, the walls of exposed spores were more susceptible to alkaline hydrolysis than those of the control spores, and some were characterized by discontinuities in the exine, anomalies in perine spine structure, abnormal intine and aperture, and occasionally, wall shedding. Our data support the involvement of oxidative cross-linking in spore-wall development, including sporopollenin polymerization or deposition, as well as a role for ROS in intine/aperture development.more » « less
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New direct numerical simulation data of a fully-developed axially rotating pipe at Re = 5300 and Re = 19, 000 is used to examine the performance of the second-moment closure elliptic blending Reynolds stress model for a range of rotation rates from N=0 to N=3. In agreement with previous studies (using alternative second-moment closure models), the turbulence suppression observed by the DNS is over-predicted. This over-prediction is greatest at Re = 5, 300 and most noticeable in the poor prediction of the ut wt turbulent shear-stress component. At N=3 the flow is completely relaminarized in contrast to the DNS that is only partly relaminarized. The accuracy of the second-moment closure model is superior to the two-equation k − ω SST model which predicts pure solid-body rotation, however, both are equally poor at the highest rotation rates. The accuracy of each model is also assessed for the initial portion of a rotating pipe where in contrast to the fully- developed rotating pipe flow the turbulent suppression is under-predicted compared to the DNS. It is clear that greater work is required to understand the root cause of the poor prediction by these second-moment closure models and further DNS and experimental work is underway to assist this effort.more » « less
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Rotating and swirling turbulence comprises an important class of flows, not only due to the complex physics that occur, but also due to their relevance to many engineering applications, such as combustion, cyclone separation, mixing, etc. In these types of flows, rotation strongly affects the characteristics and structure of turbulence. However, the underlying turbulent flow phenomena are complex and currently not well understood. The axially rotating pipe is an exemplary prototypical model problem that exhibits these complex turbulent flow physics. By examining the complex interaction of turbulent structures within rotating turbulent pipe flow, insight can be gained into the behavior of rotating flows relevant to engineering applications. Direct numerical simulations are conducted at a bulk Reynolds number up to Re_D = 19,000 with rotation numbers ranging from N = 0 to 3. Coherence analysis, including Proper Orthogonal Decomposition and Dynamic Mode Decomposition, are used to identify the relevant (highest energy) modes of the flow. Studying the influence of these modes on turbulent statistics (i.e. mean statistics, Reynolds stresses, turbulent kinetic energy, and turbulent kinetic energy budgets) allow for a deeper understanding of the effects of coherent turbulent flow structures in rotating flows.more » « less
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Highly-resolved, direct numerical simulations of turbulent channel flows with sub- Kolmogorov grid resolution are performed to investigate the characteristics of wall-bounded turbulent flows in the presence of sinusoidal wall waviness. The wall waviness serves as a simplified model to study the effects of well-defined geometric parameters of roughness on the characteristics of wall-bounded turbulent flows. In this study, a two-dimensional wave profile with steepness ranging from 0.06 to 0.25 and wave amplitudes ranging from 9 to 36 wall units were considered. For the smooth and wavy-wall simulations, the Reynolds number based on the friction velocity was kept constant. To study the effects of wave amplitude and wavelength on turbulence, two-dimensional time and spanwise averaged distributions of the mean flow, turbulent kinetic energy, and Reynolds stresses as well as turbulent kinetic energy production and dissipation are examined. Furthermore, in order to provide a more direct comparison with the smooth-wall turbulent channel flow one-dimensional pro- files of these quantities are computed by averaging them over one wavelength of the wave profile. A strong effect of the wall-waviness and, in particular, the wave amplitude and wavelength on the characteristics of the turbulence was obtained. Wall waviness mainly affected the inner flow region while all recorded turbulent statistics collapsed in the outer flow region. Significant reductions in turbulent kinetic energy, production and dissipation were obtained with increasing wave amplitudes when reported in inner scale. While production is lower for all wavy wall cases considered here in comparison to the smooth wall, reducing the wavelength caused an increase in production and a decrease in dissipation.more » « less
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