von Willebrand factor (VWF), a protein present in our circulatory system, is necessary to stop bleeding under high shear-stress conditions as found in small blood vessels. The results presented here help unravel how an increase in hydrodynamic shear stress activates VWF's adhesion potential, leading to the counterintuitive phenomena of enhanced adsorption rate under strong shear conditions. Using a microfluidic device, we were able to mimic a wide range of bloodflow conditions and directly visualize the conformational dynamics of this protein under shear flow. In particular, we find that VWF displays a reversible globule-stretch transition at a critical shear rate ␥ crit in the absence of any adsorbing surface. Computer simulations reproduce this sharp transition and identify the large size of VWF's repeating units as one of the keys for this unique hydrodynamic activation. In the presence of an adsorbing collagen substrate, we find a large increase in the protein adsorption at the same critical shear rate, suggesting that the globule unfolding in bulk triggers the surface adsorption in the case of a collagen substrate, which provides a sufficient density of binding sites. Monitoring the adsorption process of multiple VWF fibers, we were able to follow the formation of an immobilized network that constitutes a ''sticky'' grid necessary for blood platelet adhesion under high shear flow. Because areas of high shear stress coincide with a higher chance for vessel wall damage by mechanical forces, we identified the shear-induced increase in the binding probability of VWF as an effective self-regulating repair mechanism of our microvascular system. blood flow ͉ mechanical activated proteins ͉ polymer physics O ur circulatory system is exposed to an amazingly wide span of shear rates, ranging from 1 to 10 5 s Ϫ1 (1). This range calls for adhesion mechanisms during blood clotting that are different for different regimes of shear rates. At small shear rates, which are found in rather wide vessels, objects such as vesicles or cells bind to vessel walls once the contact area and the adhesion strength is large enough (2). At high shear rates, which are found in small arteries, hydrodynamic lift forces inhibit formation of a sufficiently large contact area, which makes adsorption of soft objects from the blood onto vessel walls increasingly difficult. In fact, theoretical analyses predict that adhering compact and soft objects (such as vesicles or platelets) will roll and detach from a surface at a particular shear stress and, more importantly, will remain unbound if the shear rate is increased further (3, 4). Experiments on leukocyte adhesion confirmed these theoretical predictions quite nicely (2). However, the experimental finding for blood platelet adhesion in small arteries contradicts this scenario: von Willibrand factor (VWF)-mediated platelet adhesion, which is necessary to stop bleeding in small vessels, is strongly enhanced under high shear-flow conditions (5). Therefore, this example of shear-induced adsorption m...
The measurement and simulation of water vapor isotopic composition has matured rapidly over the last decade, with long‐term data sets and comprehensive modeling capabilities now available. Theories for water vapor isotopic composition have been developed by extending the theories that have been used for the isotopic composition of precipitation to include a more nuanced understanding of evaporation, large‐scale mixing, deep convection, and kinetic fractionation. The technologies for in situ and remote sensing measurements of water vapor isotopic composition have developed especially rapidly over the last decade, with discrete water vapor sampling methods, based on mass spectroscopy, giving way to laser spectroscopic methods and satellite‐ and ground‐based infrared absorption techniques. The simulation of water vapor isotopic composition has evolved from General Circulation Model (GCM) methods for simulating precipitation isotopic composition to sophisticated isotope‐enabled microphysics schemes using higher‐order moments for water and ice size distributions. The incorporation of isotopes into GCMs has enabled more detailed diagnostics of the water cycle and has led to improvements in its simulation. The combination of improved measurement and modeling of water vapor isotopic composition opens the door to new advances in our understanding of the atmospheric water cycle, in processes ranging from the marine boundary layer, through deep convection and tropospheric mixing, and into the water cycle of the stratosphere. Finally, studies of the processes governing modern water vapor isotopic composition provide an improved framework for the interpretation of paleoclimate proxy records of the hydrological cycle.
The hydrological cycle and its response to environmental variability such as temperature changes is of prime importance for climate reconstruction and prediction. We retrieved deuterated water/water (HDO/H2O) abundances using spaceborne absorption spectroscopy, providing an almost global perspective on the near-surface distribution of water vapor isotopologs. We observed an unexpectedly high HDO/H2O seasonality in the inner Sahel region, pointing to a strong isotopic depletion in the subsiding branch of the Hadley circulation and its misrepresentation in general circulation models. An extension of the analysis at high latitudes using ground-based observations of deltaD and a model study shows that dynamic processes can entirely compensate for temperature effects on the isotopic composition of precipitation.
The behavior of a single collapsed polymer under shear flow is examined using hydrodynamic simulations and scaling arguments. Below a threshold shear rate gamma[.]{*}, the chain remains collapsed and only deforms slightly, while above gamma[.]{*} the globule exhibits unfolding/refolding cycles. Hydrodynamics are crucial: In the free draining case, gamma[.]{*} scales with the globule radius R as gamma[.]{*} approximately R{-1}, while in the presence of hydrodynamic interactions gamma[.]{*} approximately R. Experiments on the globular von Willebrand protein confirm the presence of an unfolding transition at a well-defined critical shear rate.
[1] The goal of this study is to determine how H 2 O and HDO measurements in water vapor can be used to detect and diagnose biases in the representation of processes controlling tropospheric humidity in atmospheric general circulation models (GCMs). We analyze a large number of isotopic data sets (four satellite, sixteen ground-based remote-sensing, five surface in situ and three aircraft data sets) that are sensitive to different altitudes throughout the free troposphere. Despite significant differences between data sets, we identify some observed HDO/H 2 O characteristics that are robust across data sets and that can be used to evaluate models. We evaluate the isotopic GCM LMDZ, accounting for the effects of spatiotemporal sampling and instrument sensitivity. We find that LMDZ reproduces the spatial patterns in the lower and mid troposphere remarkably well. However, it underestimates the amplitude of seasonal variations in isotopic composition at all levels in the subtropics and in midlatitudes, and this bias is consistent across all data sets. LMDZ also underestimates the observed meridional isotopic gradient and the contrast between dry and convective tropical regions compared to satellite data sets. Comparison with six other isotope-enabled GCMs from the SWING2 project shows that biases exhibited by LMDZ are common to all models. The SWING2 GCMs show a very large spread in isotopic behavior that is not obviously related to that of humidity, suggesting water vapor isotopic measurements could be used to expose model shortcomings. In a companion paper, the isotopic differences between models are interpreted in terms of biases in the representation of processes controlling humidity.Citation: Risi, C., et al. (2012), Process-evaluation of tropospheric humidity simulated by general circulation models using water vapor isotopologues: 1. Comparison between models and observations,
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