Macro- to Microscale Strain Transfer in Fibrous Tissues is Heterogeneous and Tissue-Specific

AbstractMechanical deformation applied at the joint or tissue level is transmitted through the macroscale extracellular matrix to the microscale local matrix, where it is transduced to cells within these tissues and modulates tissue growth, maintenance, and repair. The objective of this study was to investigate how applied tissue strain is transferred through the local matrix to the cell and nucleus in meniscus, tendon, and the annulus fibrosus, as well as in stem cell-seeded scaffolds engineered to reproduce the organized microstructure of these native tissues. To carry out this study, we developed a custom confocal microscope-mounted tensile testing device and simultaneously monitored strain across multiple length scales. Results showed that mean strain was heterogeneous and significantly attenuated, but coordinated, at the local matrix level in native tissues (35–70% strain attenuation). Conversely, freshly seeded scaffolds exhibited very direct and uniform strain transfer from the tissue to the local matrix level (15–25% strain attenuation). In addition, strain transfer from local matrix to cells and nuclei was dependent on fiber orientation and tissue type. Histological analysis suggested that different domains exist within these fibrous tissues, with most of the tissue being fibrous, characterized by an aligned collagen structure and elongated cells, and other regions being proteoglycan (PG)-rich, characterized by a dense accumulation of PGs and rounder cells. In meniscus, the observed heterogeneity in strain transfer correlated strongly with cellular morphology, where rounder cells located in PG-rich microdomains were shielded from deformation, while elongated cells in fibrous microdomains deformed readily. Collectively, these findings suggest that different tissues utilize distinct strain-attenuating mechanisms according to their unique structure and cellular phenotype, and these differences likely alter the local biologic response of such tissues and constructs in response to mechanical perturbation

Early Planet Formation in Embedded Disks (eDisk) VI: Kinematic Structures around the Very Low Mass Protostar IRAS 16253-2429

Precise estimates of protostellar masses are crucial to characterize the formation of stars of low masses down to brown-dwarfs (BDs; M* < 0.08 Msun). The most accurate estimation of protostellar mass uses the Keplerian rotation in the circumstellar disk around the protostar. To apply the Keplerian rotation method to a protostar at the low-mass end, we have observed the Class 0 protostar IRAS 16253-2429 using the Atacama Large Millimeter/submillimeter Array (ALMA) in the 1.3 mm continuum at an angular resolution of 0.07" (10 au), and in the 12CO, C18O, 13CO (J=2-1), and SO (J_N = 6_5-5_4) molecular lines, as part of the ALMA Large Program Early Planet Formation in Embedded Disks (eDisk). The continuum emission traces a non-axisymmetric, disk-like structure perpendicular to the associated 12CO outflow. The position-velocity (PV) diagrams in the C18O and 13CO lines can be interpreted as infalling and rotating motions. In contrast, the PV diagram along the major axis of the disk-like structure in the 12CO line allows us to identify Keplerian rotation. The central stellar mass and the disk radius are estimated to be ~0.12-0.17 Msun and ~13-19 au, respectively. The SO line suggests the existence of an accretion shock at a ring (r~28 au) surrounding the disk and a streamer from the eastern side of the envelope. IRAS 16253-2429 is not a proto-BD but has a central stellar mass close to the BD mass regime, and our results provide a typical picture of such very low-mass protostars.Comment: 41 pages, 14 figure

Early Planet Formation in Embedded Disks (eDisk) V: Possible Annular Substructure in a Circumstellar Disk in the Ced110 IRS4 System

We have observed the Class 0/I protostellar system Ced110 IRS4 at an angular resolution of $0.05''$ ($\sim$10 au) as a part of the ALMA large program; Early Planet Formation in the Embedded Disks (eDisk). The 1.3 mm dust continuum emission reveals that Ced110 IRS4 is a binary system with a projected separation of $\sim$250 au. The continuum emissions associated with the main source and its companion, named Ced110 IRS4A and IRS4B respectively, exhibit disk-like shapes and likely arise from dust disks around the protostars. The continuum emission of Ced110 IRS4A has a radius of $\sim$110 au ($\sim0.6''$), and shows bumps along its major axis with an asymmetry. The bumps can be interpreted as an shallow, ring-like structure at a radius of $\sim$40 au ($\sim0.2''$) in the continuum emission, as demonstrated from two-dimensional intensity distribution models. A rotation curve analysis on the C$^{18}$O and $^{13}$CO $J=2$-1 lines reveals the presence of a Keplerian disk within a radius of 120 au around Ced110 IRS4A, which supports the interpretation that the dust continuum emission arises from a disk. The ring-like structure in the dust continuum emission might indicate a possible, annular substructure in the surface density of the embedded disk, although the possibility that it is an apparent structure due to the optically thick continuum emission cannot be ruled out.Comment: 32 pages, 23 figures. Accepted for publication in ApJ as one of the first-look papers of the eDisk ALMA Large Progra

Early Planet Formation in Embedded Disks (eDisk). II. Limited Dust Settling and Prominent Snow Surfaces in the Edge-on Class I Disk IRAS 04302+2247

While dust disks around optically visible, Class II protostars are found to be vertically thin, when and how dust settles to the midplane are unclear. As part of the Atacama Large Millimeter/submillimeter Array (ALMA) large program, Early Planet Formation in Embedded Disks, we analyze the edge-on, embedded, Class I protostar IRAS 04302+2247, also nicknamed the ``Butterfly Star." With a resolution of 0.05" (8~au), the 1.3 mm continuum shows an asymmetry along the minor axis which is evidence of an optically thick and geometrically thick disk viewed nearly edge-on. There is no evidence of rings and gaps, which could be due to the lack of radial substructure or the highly inclined and optically thick view. With 0.1" (16~au) resolution, we resolve the 2D snow surfaces, i.e., the boundary region between freeze-out and sublimation, for $^{12}$CO $J$=2--1, $^{13}$CO $J$=2--1, C$^{18}$O $J$=2--1, $H_{2}$CO $J$=$3_{0,3}$--$2_{0,2}$, and SO $J$=$6_{5}$--$5_{4}$, and constrain the CO midplane snow line to $\sim 130$ au. We find Keplerian rotation around a protostar of $1.6 \pm 0.4 M_{\odot}$ using C$^{18}$O. Through forward ray-tracing using RADMC-3D, we find that the dust scale height is $\sim 6$ au at a radius of 100~au from the central star and is comparable to the gas pressure scale height. The results suggest that the dust of this Class~I source has yet to vertically settle significantly.Comment: 33 pages, 21 figures. Accepted for publication in ApJ as one of the first-look papers of the eDisk ALMA Large Progra

Early Planet Formation in Embedded Disks (eDisk) XII: Accretion streamers, protoplanetary disk, and outflow in the Class I source Oph IRS63

We present ALMA observations of the Class I source Oph IRS63 in the context of the Early Planet Formation in Embedded Disks (eDisk) large program. Our ALMA observations of Oph IRS63 show a myriad of protostellar features, such as a shell-like bipolar outflow (in $^{12}$CO), an extended rotating envelope structure (in $^{13}$CO), a streamer connecting the envelope to the disk (in C$^{18}$O), and several small-scale spiral structures seen towards the edge of the dust continuum (in SO). By analyzing the velocity pattern of $^{13}$CO and C$^{18}$O, we measure a protostellar mass of $\rm M_\star = 0.5 \pm 0.2$~$\rm M_\odot$ and confirm the presence of a disk rotating at almost Keplerian velocity that extends up to $\sim260$ au. These calculations also show that the gaseous disk is about four times larger than the dust disk, which could indicate dust evolution and radial drift. Furthermore, we model the C$^{18}$O streamer and SO spiral structures as features originating from an infalling rotating structure that continuously feeds the young protostellar disk. We compute an envelope-to-disk mass infall rate of $\sim 10^{-6}$~$\rm M_\odot \, yr^{-1}$ and compare it to the disk-to-star mass accretion rate of $\sim 10^{-8}$~$\rm M_\odot \, yr^{-1}$, from which we infer that the protostellar disk is in a mass build-up phase. At the current mass infall rate, we speculate that soon the disk will become too massive to be gravitationally stable.Comment: 26 pages and 17 figure

Early Planet Formation in Embedded Disks (eDisk) III: A first high-resolution view of sub-mm continuum and molecular line emission toward the Class 0 protostar L1527 IRS

Studying the physical and chemical conditions of young embedded disks is crucial to constrain the initial conditions for planet formation. Here, we present Atacama Large Millimeter/submillimeter Array (ALMA) observations of dust continuum at $\sim$0.06" (8 au) resolution and molecular line emission at $\sim$0.17" (24 au) resolution toward the Class 0 protostar L1527 IRS from the Large Program eDisk (Early Planet Formation in Embedded Disks). The continuum emission is smooth without substructures, but asymmetric along both the major and minor axes of the disk as previously observed. The detected lines of $^{12}$CO, $^{13}$CO, C$^{18}$O, H$_2$CO, c-C$_3$H$_2$, SO, SiO, and DCN trace different components of the protostellar system, with a disk wind potentially visible in $^{12}$CO. The $^{13}$CO brightness temperature and the H$_2$CO line ratio confirm that the disk is too warm for CO freeze out, with the snowline located at $\sim$350 au in the envelope. Both molecules show potential evidence of a temperature increase around the disk-envelope interface. SO seems to originate predominantly in UV-irradiated regions such as the disk surface and the outflow cavity walls rather than at the disk-envelope interface as previously suggested. Finally, the continuum asymmetry along the minor axis is consistent with the inclination derived from the large-scale (100" or 14,000 au) outflow, but opposite to that based on the molecular jet and envelope emission, suggesting a misalignment in the system. Overall, these results highlight the importance of observing multiple molecular species in multiple transitions to characterize the physical and chemical environment of young disks.Comment: 27 pages, 16 figures, 2 tables, 10 pages appendix with 12 figures. Accepted for publication in ApJ as one of the first-look papers of the eDisk ALMA Large Progra

Early Planet Formation in Embedded Disks (eDisk). IV. The Ringed and Warped Structure of the Disk around the Class I Protostar L1489 IRS

Constraining the physical and chemical structure of young embedded disks is crucial to understanding the earliest stages of planet formation. As part of the Early Planet Formation in Embedded Disks Atacama Large Millimeter/submillimeter Array Large Program, we present high spatial resolution ($\sim$0.\!\!^{\prime\prime}1 or $\sim$15 au) observations of the 1.3 mm continuum and $^{13}$CO $J=$ 2-1, C$^{18}$O $J=$ 2-1, and SO $J_N=$ $6_5$-$5_4$ molecular lines toward the disk around the Class I protostar L1489 IRS. The continuum emission shows a ring-like structure at 56 au from the central protostar and a tenuous, optically thin emission extending beyond $\sim$300 au. The $^{13}$CO emission traces the warm disk surface, while the C$^{18}$O emission originates from near the disk midplane. The coincidence of the radial emission peak of C$^{18}$O with the dust ring may indicate a gap-ring structure in the gaseous disk as well. The SO emission shows a highly complex distribution, including a compact, prominent component at $\lesssim$30 au, which is likely to originate from thermally sublimated SO molecules. The compact SO emission also shows a velocity gradient along a slightly ($\sim15^\circ$) tilted direction with respect to the major axis of the dust disk, which we interpret as an inner warped disk in addition to the warp around $\sim$200 au suggested by previous work. These warped structures may be formed by a planet or companion with an inclined orbit, or by a gradual change in the angular momentum axis during gas infall.Comment: 24 pages, 12 figures. Accepted for publication in The Astrophysical Journal as one of the first-look papers of the eDisk ALMA Large Progra

Early Planet Formation in Embedded Disks (eDisk). VII. Keplerian Disk, Disk Substructure, and Accretion Streamers in the Class 0 Protostar IRAS 16544-1604 in CB 68

We present observations of the Class 0 protostar IRAS 16544-1604 in CB 68 from the ''Early Planet Formation in Embedded Disks (eDisk)'' ALMA Large program. The ALMA observations target continuum and lines at 1.3-mm with an angular resolution of $\sim$5 au. The continuum image reveals a dusty protostellar disk with a radius of $\sim$30 au seen close to edge-on, and asymmetric structures both along the major and minor axes. While the asymmetry along the minor axis can be interpreted as the effect of the dust flaring, the asymmetry along the major axis comes from a real non-axisymmetric structure. The C$^{18}$O image cubes clearly show the gas in the disk that follows a Keplerian rotation pattern around a $\sim$0.14 $M_{\odot}$ central protostar. Furthermore, there are $\sim$1500 au-scale streamer-like features of gas connecting from North-East, North-North-West, and North-West to the disk, as well as the bending outflow as seen in the $^{12}$CO (2-1) emission. At the apparent landing point of NE streamer, there are SO (6$_5$-5$_4$) and SiO (5-4) emission detected. The spatial and velocity structure of NE streamer can be interpreted as a free-falling gas with a conserved specific angular momentum, and the detection of the SO and SiO emission at the tip of the streamer implies presence of accretion shocks. Our eDisk observations have unveiled that the Class 0 protostar in CB 68 has a Keplerian rotating disk with flaring and non-axisymmetric structure associated with accretion streamers and outflows.Comment: 30 pages, 24 figures, accepted for publication in The Astrophysical Journal as one of the first-look papers of the eDisk ALMA Large Progra

Early Planet Formation in Embedded Disks (eDisk). I. Overview of the Program and First Results

We present an overview of the Large Program, ``Early Planet Formation in Embedded Disks (eDisk)'', conducted with the Atacama Large Millimeter/submillimeter Array (ALMA). The ubiquitous detections of substructures, particularly rings and gaps, in protoplanetary disks around T Tauri stars raise the possibility that at least some planet formation may have already started during the embedded stages of star formation. In order to address exactly how and when planet formation is initiated, the program focuses on searching for substructures in disks around 12 Class 0 and 7 Class I protostars in nearby ($<$200 pc) star-forming regions through 1.3 mm continuum observations at a resolution of $\sim7$ au (0.04"). The initial results show that the continuum emission, mostly arising from dust disks around the sample protostars, has relatively few distinctive substructures, such as rings and spirals, in marked contrast to Class II disks. The dramatic difference may suggest that substructures quickly develop in disks when the systems evolve from protostars to Class II sources or alternatively that high optical depth of the continuum emission could obscure internal structures. Kinematic information obtained through CO isotopologue lines and other lines reveals the presence of Keplerian disks around protostars, providing us with crucial physical parameters, in particular, the dynamical mass of the central protostars. We describe the background of the eDisk program, the sample selection and their ALMA observations, the data reduction, and also highlight representative first-look results.Comment: This is a publication of a series of eDisk ALMA large program first-look paper

Mechanically Induced Chromatin Condensation Requires Cellular Contractility in Mesenchymal Stem Cells

This work was supported by the National Institutes of Health (R01 AR056624, R01 EB02425, T32 AR007132, and P30 AR050950). Additional support was provided by a Montague Research Award from the Perelman School of Medicine and a University of Pennsylvania University Research Foundation Award