Lamellar Bone is an Incremental Tissue Reconciling Enamel Rhythms, Body Size, and Organismal Life History. Our aim has been to conduct a series of tests to determine if lamellar bone is an incremental tissue having relevance to interpretations of organismal life history, as has been widely recognized for enamel. The first test, using modern and fossil primates and proboscideans, was designed to test the relationship between enamel striae of Retzius repeat intervals and body mass. The second test, employing primates and a few nonprimate mammals, examined the relationship between rates of lamellar bone growth and body mass, where osteocyte density was used as a surrogate for rates of bone cell proliferation in lamellar bone. Having established these relationships with body mass, the conditions were set for testing the correspondence between enamel striae of Retzius and lamellar bone development. Striae of Retzius Repeat Interval Correlates with Body Mass. Our analysis of the relationship between striae of Retzius repeat interval and body mass was performed on the largest and most taxonomically diverse sample of mammalian species to date. Significant correlations were found between these two variables, which hold true not only for the order Primates generally, including fossil hominins, but also for members of the Proboscidea, which include insular- dwarf forms. There is thus a clear association between mammalian long- period tooth growth rhythms and body mass, with a noted exception in one group of primates, the strepsirrhines (subfossil giant lemurs) having very low repeat intervals for their body mass [2. When this is considered in light of our results on the relationship between bone and tooth development, this interesting exception may be explained (see below). In addition to the interspecific comparisons examined here, the relationships between striae of Retzius repeat interval and body mass may also help to explain long- period variability within a species. Repeat intervals in Homo sapiens are highly variable, but modern humans also have an unusually high body mass variability for a primate. Preliminary research by us on six humans of known life history provides a statistically significant negative relationship between striae of Retzius repeat interval and body height (weight was extremely variable in the sample), suggesting that long period rhythms influence body size in humans (data not shown). Interspecifically, the relationship between repeat interval and body mass is positive, indicating that body mass increases are accomplished by slowing developmental rates over longer growth periods; i. Within a species and life history gamut, such as suggested by our preliminary research on humans, increases in body mass are at least partly accomplished by increasing the developmental rate within the same growth period and life history; i. Long- period rhythm variation relative to body mass thus offers a potential, if not partial, solution to questions of life history variation, while remaining cognizant of other genetic factors responsible for growth factors operating within what are otherwise restricted genetic/historical programs [2. Osteocyte Lacuna Density Correlates with Body Mass. Adobe Camera Raw F R Cs2 Molecular CompoundBased on the observed relationship between striae of Retzius repeat interval and body mass, and our appreciation for the scaling of bone mass to body mass, we advanced the working hypothesis that proliferation rates of osteoblasts, which become embedded in bone as osteocytes, vary with body mass. Osteocyte density is of special significance in this connection because, while osteoblast fates and functions are complex [3. Osteocyte lacuna densities, as they reflect the rate of osteoblast proliferation, transformation, and incorporation into bone as osteocytes during growth, should be higher in organisms with rapid growth, with a small body mass, and whose osteoblast proliferation rates would lead to higher osteocyte lacuna densities; i. No analysis has been performed to examine such a relationship within a mammalian order. We thus opted to test this hypothesis with a selection of very small to large primate and nonprimate mammal midshaft femurs (Table 4) and found that, indeed, osteocyte density is inversely proportional to body mass (Fig. 4). We ignored osteocyte apoptosis for the main reason that our interest in the development of bone mass as it relates to body mass concerns only the rate of osteoblast proliferation and the production of bone matrix by actively secreting cells. While osteocyte apoptosis most certainly occurs in some small percentage of cells, this is not a principal concern for analyses of the rate at which osteoblasts becomes embedded in the bone matrix; our interest here is independent, at least relatively so as far as we know presently, of the living bone cell population. Mullender and colleagues [3. The well- recognized reduction in the capacity for cell proliferation during postreproductive aging and disease notwithstanding [3. Though the sample is small and the analysis preliminary, it is interesting that the relationship between osteocyte lacuna density and body mass in the modern human sample is positive. Interspecifically, the relationship is negative, indicating that body mass increases are accomplished by slowing developmental rates over longer growth periods; i. However, within the human species, body mass increases are accomplished by increasing the developmental rate. These inter- and intraspecific interpretations match those described for the relationship between repeat interval and body mass above. Striae of Retzius Repeat Interval Correlates with Lamellar Bone Formation Rate. The bone lamellae of all mammals are strikingly similar in character, typical widths falling in the range of 3–6 μm, though thinner and much wider widths have been reported (to date there is no formal comparative mammalian study of lamellar widths). The lamellar design is one commonly associated with incremental patterns comprised of a layered structure, such as that represented by enamel cross striations, each layer being developed during one cycle of growth [2. The temporal association of enamel formation being known, it was our aim to examine lamellar formation rates in mammals for which the times of administration of the labels were known or could be inferred (see Materials and Methods and Table 5). In each case we found that one lamella is formed in the period matching the species- specific striae of Retzius repeat interval (e. Fig. 5, Table 5). For example, in the pigtailed macaque (Fig. 5), one vital label to the next was observed to contain between three and four lamellae, followed by four or five lamellae from the last label to the end of growth, which is consistent with a lamellar formation rate of 4 days/lamella and the striae of Retzius repeat interval for this taxon [8]. The lamellar structure in Fig. 5 contains some anisotropy, however, which is increased in the lower portion of the image. Concerning this anisotropy, there are two points to note. · Initially thought to be a means for cells to eliminate waste, secreted extracellular vesicles, known as exosomes, are now understood to mediate numerous. Tabtight professional, free when you need it, VPN service. First, where one lamella is observed in the upper portion of this image, two may arise below. Second, the fluorescence image derives from a single confocal plane overlain onto a circularly polarized light image of the full section thickness, thus vital labels are observed to waver slightly around lamellae; i. Adobe Camera Raw F R Cs2 Molecular ShapeLamellar Bone is an Incremental Tissue Reconciling Enamel Rhythms, Body Size, and Organismal Life History. · Functional Characterization of MAT1-1-Specific Mating-Type Genes in the Homothallic Ascomycete Sordaria macrospora Provides New Insights into Essential and. Both of these phenomena are explained by the anisotropy discussed under Materials and Methods, which is informative and helps us to appreciate the three- dimensional complexity of forming bone surfaces. In both humans and sheep it was also found that the lamellar formation rate was identical in both primary lamellar bone laid down during growth and secondary remodeled osteonal bone laid down during bone maintenance. Evidence for primary and secondary lamellar bone formation combined demonstrates that lamellar bone is an incremental tissue with a consistent time- dependent developmental morphology, similar to that described for enamel. Long- Period Rhythms and Organismal Life History. Because the bone organ provides mechanical support for body mass, developmental rates for body mass should reflect developmental rates for bone mass. We thus hypothesize that the metabolic rhythm affecting osteoblast behavior and lamellar bone formation sets the pace of skeletal development; the effect this rhythm has on ameloblast function and enamel formation is secondary. A Polyethylene Glycol- Based Method for Enrichment of Extracellular Vesicles. Abstract. Initially thought to be a means for cells to eliminate waste, secreted extracellular vesicles, known as exosomes, are now understood to mediate numerous healthy and pathological processes. Though abundant in biological fluids, purifying exosomes has been challenging because their biophysical properties overlap with other secreted cell products. Easy- to- use commercial kits for harvesting exosomes are now widely used, but the relative low- purity and high- cost of the preparations restricts their utility. Here we describe a method for purifying exosomes and other extracellular vesicles by adapting methods for isolating viruses using polyethylene glycol. This technique, called Extra. PEG, enriches exosomes from large volumes of media rapidly and inexpensively using low- speed centrifugation, followed by a single small- volume ultracentrifugation purification step. Total protein and RNA harvested from vesicles is sufficient in quantity and quality for proteomics and sequencing analyses, demonstrating the utility of this method for biomarker discovery and diagnostics. Additionally, confocal microscopy studies suggest that the biological activity of vesicles is not impaired. The Extra. PEG method can be easily adapted to enrich for different vesicle populations, or as an efficient precursor to subsequent purification techniques, providing a means to harvest exosomes from many different biological fluids and for a wide variety of purposes. Exosomes are virus- sized membranous sacs released from cells. They contain protein and nucleic acid cargo that mediate intercellular communication. Exosomes are present in diverse biological fluids and contain molecular signatures of their progenitor cells. Additionally, the discovery that exosomes participate in pathological conditions has generated tremendous interest in uncovering their functions, and efforts to discover biomarkers in exosomes for diagnostic or prognostic purposes have intensified. Exosomes are distinguished from other extracellular vesicles by size, content, and location of their biogenesis. Whereas other extracellular vesicles, like microvesicles, bud off from the cell surface, exosomes are generally regarded as vesicles that originate at internal endosomal membranes of multivesicular bodies (MVB), and are released into the extracellular milieu following fusion of the MVB with the plasma membrane. The most commonly used exosome enrichment procedure is differential centrifugation (DC): a series of centrifugation steps of increasing centrifugal force (typically up to 1. Exosome isolation is complicated by the fact that vesicle subpopulations are not thoroughly defined and may overlap in size and density. Furthermore, there is ongoing debate about the presence of exosome subpopulations with unique cargo and mechanisms of formation. This makes achieving truly pure vesicle preparations empirically impossible. Adding to this problem is the great expense and handling time required for enrichment and purification, especially for proteomics experiments that require concentrating large volumes of media for extracting vesicle protein. Recently, user- friendly but expensive commercial precipitation reagents like Exo. Quick. TM from Systems Biosciences, and Total Exosome Isolation. TM from Life Technologies have become popular. Despite widespread use of these reagents, very little has been reported about the quality and quantity of the exosomes harvested compared to other methods. Exo. Quick (patent number: US2. A1) and Total Exosome Isolation (TEI; US2. A1) reagents contain volume- excluding polymers (e. However, simple solutions of polyethylene glycol (PEG) have been used for over fifty years to concentrate and purify viruses. Because exosomes and virus particles have similar biophysical properties, we hypothesized that a PEG- based method used for virus isolation could be modified to enrich and purify exosomes, providing an inexpensive and efficient alternative to commercially available products and DC. We developed and optimized a protocol for exosome enrichment based on Mahy and Kangro’s compendium on the use of PEG for virus enrichment. To assess our adapted protocol, we first attempted to enrich vesicles from healthy cell cultures. Then, we aimed to evaluate the practical utility of this method by comparing it to the most commonly used methods for enriching vesicles and exosomes. Finally, we tested the versatility of this method by investigating the biological activity of enriched vesicles, the ability to collect from them intact protein and RNA, and by evaluating vesicle yields from a diversity of biological fluids. Overall, we sought a rapid, inexpensive, and adaptable means of harvesting large amounts of exosomes without sacrificing the purity of the preparation. Results. Polyethylene glycol treatment of cell culture medium enriches exosomes. To demonstrate that a simple adaptation of the virus enrichment method would enrich exosomes, PEG solutions were added to culture media at concentrations between 5 and 1. The following day, samples were centrifuged for 1 hr at maximum speed in a standard tabletop centrifuge (to maximize ease of processing large volumes of media). The resulting pellets were suspended and particles were characterized using a nanoparticle tracker. Regression analysis indicated a strong positive association between the concentration of PEG and particle numbers recovered (p < 0. R- squared = 0. 9. Fig. 1a). All experimental conditions yielded exosome- sized particles, and particle size (mean and mode) did not differ significantly between any of the groups (Fig. There was, however, a negative association between mode particle size and PEG concentration (p < 0. R- squared = 0. 3. This suggests that higher PEG concentrations foster precipitation of smaller extracellular vesicles. Alternatively, these results could represent increased precipitation of off- target protein aggregates. As serum proteins are common contaminants in vesicle preparations from cell culture, the purity of the preparations was also assessed by calculating the ratio of particle numbers harvested per total protein harvested (particle number in 1 m. L medium per microgram quantity of protein in the same volume medium; Fig. Purity of the preparations peaked with PEG concentrations of between 6 and 8% (with no statistical difference between those sample groups). At 9% PEG, there was a statistically significant drop in purity from the maximum (p = 0. Polyethylene glycol enriches vesicles containing exosome proteins. The presence of targeted vesicles was confirmed by western blot analyses probing for the markers of exosomes, TSG1. CD6. 3, HSC7. 0, and ALIX1. Each lane of the gel contained an equal volume of lysate from a treatment group (Fig. Samples using higher PEG concentration produced greater signal intensity, corroborating the nanoparticle tracking results. The same samples were then reevaluated, but this time an equal microgram quantity of each lysate was loaded into the gel to assess purity. Precipitation of off- target proteins was expected to increase with increasing concentrations of PEG, thus reducing the amount of exosome protein loaded into the gel. As expected, for most markers signal strength peaked in groups treated with 7 to 8% PEG (Fig. The pattern of HSC7. HSC7. 0 may be enriched in smaller vesicle populations or protein complexes that are preferentially isolated using higher PEG concentrations. Finally, qualitative assessment of the proteins was conducted by staining the nitrocellulose membrane blot with Ponceau S solution (Fig. Similar banding patterns appeared in lanes loaded with FBS alone, suggesting serum protein predominated amongst the harvested protein (see also Fig. As PEG concentration increased, so did the quantity and composition of serum proteins that co- precipitated with vesicle proteins. Consistent with the western analysis, at 7 to 8% fewer unique serum proteins were harvested. PEG- based vesicle enrichment is comparable or superior to standard and commercial methods. Having confirmed that the enriched particles contained exosome protein, techniques were explored for improving yield and purity, and these methods were compared to the commercially available Exo. Quick and Total Exosome Isolation methods. As the 8% PEG solution yielded high amounts of particles without significantly sacrificing purity (Fig.
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In his 1. 68. 7 theory, Isaac Newton postulated space as an infinite and unalterable physical structure existing before, within, and around all objects while their states and relations unfold at a constant pace everywhere, thus absolute space and time.Inferring that all objects bearing mass approach at a constant rate, but collide by impact proportional to their masses, Newton inferred that matter exhibits an attractive force.His law of universal gravitation mathematically stated it to span the entire universe instantly (despite absolute time), or, if not actually a force, to be instant interaction among all objects (despite absolute space.) As conventionally interpreted, Newton's theory of motion modelled a central force without a communicating medium. Thus Newton's theory violated the first principle of mechanical philosophy, as stated by Descartes, No action at a distance. Conversely, during the 1. Michael Faraday inferred a field filling space and transmitting that force. Faraday conjectured that ultimately, all forces unified into one. In the early 1. 87. James Clerk Maxwell unified electricity and magnetism as effects of an electromagnetic field whose third consequence was light, travelling at constant speed in a vacuum. The electromagnetic field theory contradicted predictions of Newton's theory of motion, unless physical states of the luminiferous aether—presumed to fill all space whether within matter or in a vacuum and to manifest the electromagnetic field—aligned all phenomena and thereby held valid the Newtonian principle relativity or invariance. The exchange of bosons always carries energy and momentum between the fermions, thereby changing their speed and direction. The exchange may also transport a charge between the fermions, changing the charges of the fermions in the process (e. Since bosons carry one unit of angular momentum, the fermion's spin direction will flip from +? Planck's constant). Because an interaction results in fermions attracting and repelling each other, an older term for "interaction" is force. According to the present understanding, there are four fundamental interactions or forces: gravitation, electromagnetism, the weak interaction, and the strong interaction. Their magnitude and behaviour vary greatly, as described in the table below. Modern physics attempts to explain every observed physical phenomenon by these fundamental interactions. Moreover, reducing the number of different interaction types is seen as desirable. Two cases in point are the unification of. Electric and magnetic force into electromagnetism. The electromagnetic interaction and the weak interaction into the electroweak interaction; see below. Both magnitude ("relative strength") and "range", as given in the table, are meaningful only within a rather complex theoretical framework. 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