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This endpoint is enabled and enabled for proxy by default. The FederationMetadata. Once the federation trust is created between partners, the Federation Service holds the Federation Metadata endpoint as a property of its partners, and uses the endpoint to periodically check for updates from the partner.
For example, if an Identity Provider gets a new token-signing certificate, the public key portion of that certificate is published as part of its Federation Metadata. All Relying Parties who partner with this IdP will automatically be able to validate the digital signature on tokens issued by the IdP because the RP has refreshed the Federation Metadata via the endpoint.
The Federation Metadata. XML publishes information such as the public key portion of a token signing certificate and the public key of the Encryption Certificate. What we can do is creating a schedule process which:. You can create the source with the following line as an Administrator of the server:. Signing Certificate. Encryption Certificate. As part of my Mix and Match series , we went through concepts and terminologies of the Identity metasystem, understood how all the moving parts operates across organizational boundaries.
We discussed the certificates involvement in AD FS and how I can use PowerShell to create a custom monitor workload and a proper logging which can trigger further automation. I hope you have enjoyed and that this can help you if you land on this page. Hi everyone, Robert Smith here to talk to you today a bit about crash dump configurations and options. With the wide-spread adoption of virtualization, large database servers, and other systems that may have a large amount or RAM, pre-configuring the systems for the optimal capturing of debugging information can be vital in debugging and other efforts.
Ideally a stop error or system hang never happens. But in the event something happens, having the system configured optimally the first time can reduce time to root cause determination. The information in this article applies the same to physical or virtual computing devices. You can apply this information to a Hyper-V host, or to a Hyper-V guest.
You can apply this information to a Windows operating system running as a guest in a third-party hypervisor. If you have never gone through this process, or have never reviewed the knowledge base article on configuring your machine for a kernel or complete memory dump , I highly suggest going through the article along with this blog.
When a windows system encounters an unexpected situation that could lead to data corruption, the Windows kernel will implement code called KeBugCheckEx to halt the system and save the contents of memory, to the extent possible, for later debugging analysis. The problem arises as a result of large memory systems, that are handling large workloads. Even if you have a very large memory device, Windows can save just kernel-mode memory space, which usually results in a reasonably sized memory dump file.
But with the advent of bit operating systems, very large virtual and physical address spaces, even just the kernel-mode memory output could result in a very large memory dump file. When the Windows kernel implements KeBugCheckEx execution of all other running code is halted, then some or all of the contents of physical RAM is copied to the paging file. On the next restart, Windows checks a flag in the paging file that tells Windows that there is debugging information in the paging file. Please see KB for more information on this hotfix.
Herein lies the problem. One of the Recovery options is memory dump file type. There are a number of memory. For reference, here are the types of memory dump files that can be configured in Recovery options:. Anything larger would be impractical. For one, the memory dump file itself consumes a great deal of disk space, which can be at a premium.
Second, moving the memory dump file from the server to another location, including transferring over a network can take considerable time.
The file can be compressed but that also takes free disk space during compression. The memory dump files usually compress very well, and it is recommended to compress before copying externally or sending to Microsoft for analysis.
On systems with more than about 32 GB of RAM, the only feasible memory dump types are kernel, automatic, and active where applicable. Kernel and automatic are the same, the only difference is that Windows can adjust the paging file during a stop condition with the automatic type, which can allow for successfully capturing a memory dump file the first time in many conditions. A 50 GB or more file is hard to work with due to sheer size, and can be difficult or impossible to examine in debugging tools.
In many, or even most cases, the Windows default recovery options are optimal for most debugging scenarios. The purpose of this article is to convey settings that cover the few cases where more than a kernel memory dump is needed the first time. Nobody wants to hear that they need to reconfigure the computing device, wait for the problem to happen again, then get another memory dump either automatically or through a forced method.
The problem comes from the fact that the Windows has two different main areas of memory: user-mode and kernel-mode. User-mode memory is where applications and user-mode services operate. Kernel-mode is where system services and drivers operate. This explanation is extremely simplistic. More information on user-mode and kernel-mode memory can be found at this location on the Internet:.
User mode and kernel mode. What happens if we have a system with a large amount of memory, we encounter or force a crash, examine the resulting memory dump file, and determine we need user-mode address space to continue analysis? This is the scenario we did not want to encounter. We have to reconfigure the system, reboot, and wait for the abnormal condition to occur again. The secondary problem is we must have sufficient free disk space available. If we have a secondary local drive, we can redirect the memory dump file to that location, which could solve the second problem.
The first one is still having a large enough paging file. If the paging file is not large enough, or the output file location does not have enough disk space, or the process of writing the dump file is interrupted, we will not obtain a good memory dump file.
In this case we will not know until we try. Wait, we already covered this. The trick is that we have to temporarily limit the amount of physical RAM available to Windows. The numbers do not have to be exact multiples of 2. The last condition we have to meet is to ensure the output location has enough free disk space to write out the memory dump file. Once the configurations have been set, restart the system and then either start the issue reproduction efforts, or wait for the abnormal conditions to occur through the normal course of operation.
Note that with reduced RAM, there ability to serve workloads will be greatly reduced. Once the debugging information has been obtained, the previous settings can be reversed to put the system back into normal operation. This is a lot of effort to go through and is certainly not automatic. But in the case where user-mode memory is needed, this could be the only option.
Figure 1: System Configuration Tool. Figure 2: Maximum memory boot configuration. Figure 3: Maximum memory set to 16 GB. With a reduced amount of physical RAM, there may now be sufficient disk space available to capture a complete memory dump file. In the majority of cases, a bugcheck in a virtual machine results in the successful collection of a memory dump file. The common problem with virtual machines is disk space required for a memory dump file.
The default Windows configuration Automatic memory dump will result in the best possible memory dump file using the smallest amount of disk space possible.
The main factors preventing successful collection of a memory dump file are paging file size, and disk output space for the resulting memory dump file after the reboot. These drives may be presented to the VM as a local disk, that can be configured as the destination for a paging file or crashdump file.
The problem occurs in case a Windows virtual machine calls KeBugCheckEx , and the location for the Crashdump file is configured to write to a virtual disk hosted on a file share. Depending on the exact method of disk presentation, the virtual disk may not be available when needed to write to either the paging file, or the location configured to save a crashdump file. It may be necessary to change the crashdump file type to kernel to limit the size of the crashdump file.
Either that, or temporarily add a local virtual disk to the VM and then configure that drive to be the dedicated crashdump location. How to use the DedicatedDumpFile registry value to overcome space limitations on the system drive when capturing a system memory dump.
The important point is to ensure that a disk used for paging file, or for a crashdump destination drive, are available at the beginning of the operating system startup process. Virtual Desktop Infrastructure is a technology that presents a desktop to a computer user, with most of the compute requirements residing in the back-end infrastructure, as opposed to the user requiring a full-featured physical computer. Usually the VDI desktop is accessed via a kiosk device, a web browser, or an older physical computer that may otherwise be unsuitable for day-to-day computing needs.
Non-persistent VDI means that any changes to the desktop presented to the user are discarded when the user logs off. Even writes to the paging file are redirected to the write cache disk. Typically the write cache disk is sized for normal day-to-day computer use. The problem occurs that, in the event of a bugcheck, the paging file may no longer be accessible. Even if the pagefile is accessible, the location for the memory dump would ultimately be the write cache disk. Even if the pagefile on the write cache disk could save the output of the bugcheck data from memory, that data may be discarded on reboot.
Even if not, the write cache drive may not have sufficient free disk space to save the memory dump file. In the event a Windows operating system goes non-responsive, additional steps may need to be taken to capture a memory dump. Setting a registry value called CrashOnCtrlScroll provides a method to force a kernel bugcheck using a keyboard sequence. This will trigger the bugcheck code, and should result in saving a memory dump file.
A restart is required for the registry value to take effect. This situation may also help in the case of accessing a virtual computer and a right CTRL key is not available. For server-class, and possibly some high-end workstations, there is a method called Non-Maskable Interrupt NMI that can lead to a kernel bugcheck. The NMI method can often be triggered over the network using an interface card with a network connection that allows remote connection to the server over the network, even when the operating system is not running.
In the case of a virtual machine that is non-responsive, and cannot otherwise be restarted, there is a PowerShell method available.
This command can be issued to the virtual machine from the Windows hypervisor that is currently running that VM. The big challenge in the cloud computing age is accessing a non-responsive computer that is in a datacenter somewhere, and your only access method is over the network. In the case of a physical server there may be an interface card that has a network connection, that can provide console access over the network.
Other methods such as virtual machines, it can be impossible to connect to a non-responsive virtual machine over the network only. The trick though is to be able to run NotMyFault. If you know that you are going to see a non-responsive state in some amount of reasonable time, an administrator can open an elevated.
Some other methods such as starting a scheduled task, or using PSEXEC to start a process remotely probably will not work, because if the system is non-responsive, this usually includes the networking stack. Hopefully this will help you with your crash dump configurations and collecting the data you need to resolve your issues. Hello Paul Bergson back again, and I wanted to bring up another security topic. There has been a lot of work by enterprises to protect their infrastructure with patching and server hardening, but one area that is often overlooked when it comes to credential theft and that is legacy protocol retirement.
To better understand my point, American football is very fast and violent. Professional teams spend a lot of money on their quarterbacks. Quarterbacks are often the highest paid player on the team and the one who guides the offense. There are many legendary offensive linemen who have played the game and during their time of play they dominated the opposing defensive linemen.
Over time though, these legends begin to get injured and slow down do to natural aging. Unfortunately, I see all too often, enterprises running old protocols that have been compromised, with in the wild exploits defined, to attack these weak protocols. TLS 1. The WannaCrypt ransomware attack, worked to infect a first internal endpoint. The initial attack could have started from phishing, drive-by, etc… Once a device was compromised, it used an SMB v1 vulnerability in a worm-like attack to laterally spread internally.
A second round of attacks occurred about 1 month later named Petya, it also worked to infect an internal endpoint. Once it had a compromised device, it expanded its capabilities by not only laterally moving via the SMB vulnerability it had automated credential theft and impersonation to expand on the number devices it could compromise. This is why it is becoming so important for enterprises to retire old outdated equipment, even if it still works! The above listed services should all be scheduled for retirement since they risk the security integrity of the enterprise.
The cost to recover from a malware attack can easily exceed the costs of replacement of old equipment or services. Improvements in computer hardware and software algorithms have made this protocol vulnerable to published attacks for obtaining user credentials. As with any changes to your environment, it is recommended to test this prior to pushing into production. If there are legacy protocols in use, an enterprise does run the risk of services becoming unavailable.
To disable the use of security protocols on a device, changes need to be made within the registry. Once the changes have been made a reboot is necessary for the changes to take effect. The registry settings below are ciphers that can be configured. Note: Disabling TLS 1. Microsoft highly recommends that this protocol be disabled. KB provides the ability to disable its use, but by itself does not prevent its use.
For complete details see below. The PowerShell command above will provide details on whether or not the protocol has been installed on a device. Ralph Kyttle has written a nice Blog on how to detect, in a large scale, devices that have SMBv1 enabled. Once you have found devices with the SMBv1 protocol installed, the device should be monitored to see if it is even being used.
Open up Event Viewer and review any events that might be listed. The tool provides client and web server testing. From an enterprise perspective you will have to look at the enabled ciphers on the device via the Registry as shown above. If it is found that it is enabled, prior to disabling, Event Logs should be inspected so as to possibly not impact current applications. Hello all! Nathan Penn back again with a follow-up to Demystifying Schannel.
While finishing up the original post, I realized that having a simpler method to disable the various components of Schannel might be warranted. If you remember that article, I detailed that defining a custom cipher suite list that the system can use can be accomplished and centrally managed easily enough through a group policy administrative template. However, there is no such administrative template for you to use to disable specific Schannel components in a similar manner.
The result being, if you wanted to disable RC4 on multiple systems in an enterprise you needed to manually configure the registry key on each system, push a registry key update via some mechanism, or run a third party application and manage it. Well, to that end, I felt a solution that would allow for centralized management was a necessity, and since none existed, I created a custom group policy administrative template.
The administrative template leverages the same registry components we brought up in the original post, now just providing an intuitive GUI. For starters, the ever-important logging capability that I showcased previously, has been built-in. So, before anything gets disabled, we can enable the diagnostic logging to review and verify that we are not disabling something that is in use.
While many may be eager to start disabling components, I cannot stress the importance of reviewing the diagnostic logging to confirm what workstations, application servers, and domain controllers are using as a first step. Once we have completed that ever important review of our logs and confirmed that components are no longer in use, or required, we can start disabling.
Within each setting is the ability to Enable the policy and then selectively disable any, or all, of the underlying Schannel components. Remember, Schannel protocols, ciphers, hashing algorithms, or key exchanges are enabled and controlled solely through the configured cipher suites by default, so everything is on. To disable a component, enable the policy and then checkbox the desired component that is to be disabled. Note, that to ensure that there is always an Schannel protocol, cipher, hashing algorithm, and key exchange available to build the full cipher suite, the strongest and most current components of each category was intentionally not added.
Finally, when it comes to practical application and moving forward with these initiatives, start small. I find that workstations is the easiest place to start. Create a new group policy that you can security target to just a few workstations. Enable the logging and then review.
Then re-verify that the logs show they are only using TLS. At this point, you are ready to test disabling the other Schannel protocols. Once disabled, test to ensure the client can communicate out as before, and any client management capability that you have is still operational. If that is the case, then you may want to add a few more workstations to the group policy security target.
And only once I am satisfied that everything is working would I schedule to roll out to systems in mass. After workstations, I find that Domain Controllers are the next easy stop. With Domain Controllers, I always want them configured the identically, so feel free to leverage a pre-existing policy that is linked to the Domain Controllers OU and affects them all or create a new one.
The important part here is that I review the diagnostic logging on all the Domain Controllers before proceeding. Lastly, I target application servers grouped by the application, or service they provide. Working through each grouping just as I did with the workstations. Creating a new group policy, targeting a few systems, reviewing those systems, re-configuring applications as necessary, re-verifying, and then making changes. Both of these options will re-enable the components the next time group policy processes on the system.
To leverage the custom administrative template we need to add them to our Policy Definition store. Once added, the configuration options become available under:. Each option includes a detailed description of what can be controlled as well as URLs to additional information.
You can download the custom Schannel ADM files by clicking here! I could try to explain what the krbtgt account is, but here is a short article on the KDC and the krbtgt to take a look at:.
Both items of information are also used in tickets to identify the issuing authority. For information about name forms and addressing conventions, see RFC This provides cryptographic isolation between KDCs in different branches, which prevents a compromised RODC from issuing service tickets to resources in other branches or a hub site. The RODC does not have the krbtgt secret. Thus, when removing a compromised RODC, the domain krbtgt account is not lost.
So we asked, what changes have been made recently? In this case, the customer was unsure about what exactly happened, and these events seem to have started out of nowhere. They reported no major changes done for AD in the past 2 months and suspected that this might be an underlying problem for a long time. So, we investigated the events and when we looked at it granularly we found that the event was coming from a RODC:.
Computer: ContosoDC. Internal event: Active Directory Domain Services could not update the following object with changes received from the following source directory service. This is because an error occurred during the application of the changes to Active Directory Domain Services on the directory service. To reproduce this error in lab we followed the below steps: —. If you have a RODC in your environment, do keep this in mind.
Thanks for reading, and hope this helps! Hi there! Windows Defender Antivirus is a built-in antimalware solution that provides security and antimalware management for desktops, portable computers, and servers. This library of documentation is aimed for enterprise security administrators who are either considering deployment, or have already deployed and are wanting to manage and configure Windows Defender AV on PC endpoints in their network.
Nathan Penn and Jason McClure here to cover some PKI basics, techniques to effectively manage certificate stores, and also provide a script we developed to deal with common certificate store issue we have encountered in several enterprise environments certificate truncation due to too many installed certificate authorities. To get started we need to review some core concepts of how PKI works. Some of these certificates are local and installed on your computer, while some are installed on the remote site.
The lock lets us know that the communication between our computer and the remote site is encrypted. But why, and how do we establish that trust? Regardless of the process used by the site to get the certificate, the Certificate Chain, also called the Certification Path, is what establishes the trust relationship between the computer and the remote site and is shown below.
As you can see, the certificate chain is a hierarchal collection of certificates that leads from the certificate the site is using support. To establish the trust relationship between a computer and the remote site, the computer must have the entirety of the certificate chain installed within what is referred to as the local Certificate Store.
When this happens, a trust can be established and you get the lock icon shown above. But, if we are missing certs or they are in the incorrect location we start to see this error:.
The primary difference being that certificates loaded into the Computer store become global to all users on the computer, while certificates loaded into the User store are only accessible to the logged on user. To keep things simple, we will focus solely on the Computer store in this post. Leveraging the Certificates MMC certmgr. This tool also provides us the capability to efficiently review what certificates have been loaded, and if the certificates have been loaded into the correct location.
Trusted Root CAs are the certificate authority that establishes the top level of the hierarchy of trust. By definition this means that any certificate that belongs to a Trusted Root CA is generated, or issued, by itself. Simple stuff, right? We know about remote site certificates, the certificate chain they rely on, the local certificate store, and the difference between Root CAs and Intermediate CAs now. But what about managing it all? On individual systems that are not domain joined, managing certificates can be easily accomplished through the same local Certificates MMC shown previously.
In addition to being able to view the certificates currently loaded, the console provides the capability to import new, and delete existing certificates that are located within. Using this approach, we can ensure that all systems in the domain have the same certificates loaded and in the appropriate store. It also provides the ability to add new certificates and remove unnecessary certificates as needed.
On several occasions both of us have gone into enterprise environments experiencing authentication oddities, and after a little analysis trace the issue to an Schannel event This list has thus been truncated. On a small scale, customers that experience certificate bloat issues can leverage the Certificate MMC to deal with the issue on individual systems. Unfortunately, the ability to clear the certificate store on clients and servers on a targeted and massive scale with minimal effort does not exist.
This technique requires the scripter to identify and code in the thumbprint of every certificate that is to be purged on each system also very labor intensive. Only certificates that are being deployed to the machine from Group Policy will remain. The ability to clear the certificate store on clients and servers on a targeted and massive scale with minimal effort. This is needed to handle certificate bloat issues that can ultimately result in authentication issues.
On a small scale, customers that experience certificate bloat issues can leverage the built-in certificate MMC to deal with the issue on a system by system basis as a manual process. CertPurge then leverages the array to delete every subkey. Prior to performing any operations i.
In the event that required certificates are purged, an administrator can import the backup files and restore all purged certificates. NOTE: This is a manual process, so testing prior to implementation on a mass scale is highly recommended.
KB details the certificates that are required for the operating system to operate correctly. Removal of the certificates identified in the article may limit functionality of the operating system or may cause the computer to fail.
If a required certificate either one from the KB, or one specific to the customer environment is purged, that is not being deployed via GPO, the recommended approach is as follows. Restore certificates to an individual machine using the backup registry file,. Leveraging the Certificate MMC, export the required certificates to file,.
Update the GPO that is deploying certificates by importing the required certificates,. Rerun CertPurge on machine identified in step 1 to re-purge all certificates,. Did we mention Test? Also, we now have a method for cleaning things up things in bulk should things get out of control and you need to rebaseline systems in mass.
All KB and revision numbers are documented on Microsoft Documentation. On a device running Windows 11 or Windows 10, you can run winver in a command window. You can also use this useful Powershell script from Trevor Jones. The script will show you:. You can use various tools in the SCCM console to do so.
If you want to create collections based on Windows 10 versions, you can use our set of Operational Collections or use this query. You only need to change your version number at the end of the query. The Windows servicing information is spread across many views. If you need to build a Windows 10 report you can use these views to get your information. With time, I added more and more collections to the script.
Fast forward to today, the script now contains collections and has been downloaded more than 75 times making this PowerShell script my most downloaded contribution to the community. The collections are set to refresh on a 7 days schedule.
Once created, you can use these collections to have a quick overview of your devices. You can also use these collections to create deployment collections by using limiting collections on these ones. The script will detect if the collection has already been created. It will give a warning and create only new collections that have been added since the last time the script is run. If you are comfortable with editing scripts, you can comment out any unwanted collections using at each line of the section.
Extra hint: You can also verify if your collection has been created properly in your collections with our Configuration Manager — Collections report. Simply sort the report by the Operational folder name. If you want to add a collection to the list, feel free to contact me using our social media or use the comment section. It will be our pleasure to add it to the next version. The reason to Customize Windows Start Menu is a must for any organization to deploy a standard workstation and remove any unwanted software from it.
Sometimes Microsoft makes small changes under the hood and can hardly be tracked unless an issue comes up to flag those changes. Windows 11 which came out recently share the same mechanism as Windows 10 when it comes to the Start Menu thus, this post can be used for Windows Microsoft added the following note to the start menu layout modification documentation after the release. Following our previous posts on Windows 10 Customization and how to modify the taskbar configuration , we will detail how to configure the start menu and taskbar with the latest indication from Microsoft.
Once this is completed it can be added to your SCCM task sequence like we explain in our previous posts. Comanagement enables some interesting features like conditional access, remote actions with Intune, and provisioning using AutoPilot.
This is great to slowly phase into Intune. Microsoft provides a great diagram that explains how the workload is managed when co-management is activated. The co-management provides the ability to offload some workload to Intune. There are 3 categories of workloads :. The co-management is designed to allow administrators to Pilot to specific computers before completely offloading a workload to Intune, allowing a smooth transition.
After MDM managed. More details about switching workload to Intune on Microsoft learn. In this post, we will be looking at using SCCM dynamic queries to populate collections in our deployments.
As an SCCM administrator, you most likely had to plan out mass deployments to all your servers or workstations or even both. How did you go ahead and populate your collections? Since the introduction of SCCM , we now have a multitude of options, most notably:. Chances are, if you are deploying new software to be part of a baseline for workstations for example , you will also add it to your task sequence.
In my past life, I must admit, I really did like queries. They can be such a powerful tool to populate your collections. I always was looking for ways to pimp the usual types of queries we use. For example, we developed a fabulous list of operational collections that we can use for our day-to-day deployments. But, that stays static. What I mean by that is if your collection targets workstations, you will always target workstations minus or more of the workstations that get added as the query gets updated.
I personally like when things are a little more dynamic. If I target a deployment for workstations, I would like to see that collection drop to 50, 40, 25 or whatever the count of objects as the deployment succeeds on workstations.
We have a deployment. We want to deploy this on all our workstations. Simple right? What if we add to the same query another criteria that exclude all workstations where the Deployment ID for 7-Zip is successful? As the workstations install the software and return a success code to their management point, this query will rerun itself and should yield fewer and fewer objects.
Now, you can use this for all your deployments. But to be optimal, you need to use Package deployments and not applications. So I stated earlier, we start with a very basic package for 7-Zip. And as we typically do, this program is deployed to a collection, in this case I went very originally with Deploy 7-Zip. Nothing special with our collection the way we usually do it. My current query lists a grand total of 4 objects in my collection. You can clearly see the type of rule is set to Query.
Note: I set my updates on collections at 30 minutes. This is my personal lab. I would in no case set this for a real live production collection. Most aggressive I would typically go for would be 8 hours. Understanding WQL can be a challenge if you never played around with it.
Press Ok. As you can see in the screenshot below, my count went down by two since I already had successfully deployed it to half my test machines.
Ok, now that we have that dynamic query up and running, why not try and improve on the overall deployment technique, shall we? As you know, a program will be deployed when the Assignment schedule time is reached. If you have computers that are offline, they will receive their installation when they boot up their workstation, unless you have a maintenance window preventing it. Unless you have set a recurring schedule, it will not rerun. By having a dynamic collection as we did above, combined with a recurring schedule, you can reattempt the installation on all workstations that failed the installation without starting the process for nothing on a workstation that succeeded to install it.
As I said earlier, the goal of this post is not necessarily to replace your deployment methods. By targeting the SCCM client installation error codes, you will have a better idea of what is happening during client installation. The error codes are not an exact science, they can defer depending on the situation. For a better understanding of ccmsetup error codes, read this great post from Jason Sandys. A better SCCM client installation rate equals better overall management.
POC caster circulates the opinions of community members by using conversational representation in a broadcasting system on the Internet. We evaluated its transformation rules in two experiments.
In Experiment 1, we examined our transformation rules for conversational representation in relation to sentence length. Log in with Facebook Log in with Google. Remember me on this computer. Enter the email address you signed up with and we’ll email you a reset link.
Need an account? Click here to sign up. Download Free PDF. Siu-Tsen Shen. Related Papers. However, because of certain types of intrinsic disorder present in the otherwise well-ordered sarcomeres these images did not resolve the individual myosin heads or actin subunits. Electron tomography ET produced more informative 3D images without spatial averaging, but with a significant noise component, making direct interpretation difficult. The later development of subvolume classification and averaging provided the most detailed images of myosin heads operating in situ in the muscle.
Certain features of the actin-myosin interaction, particularly those indicating large azimuthal changes in the lever arm orientation relative to crystal structures of myosin heads, have been remarkably consistent among the many structures reported over the years.
The supply of Lethocerus is seasonal. Unless muscle tissue can be dissected and used immediately, it is generally glycerinated [ 35 ]. Muscle fibers and even individual myofibrils are too thick for examination in an EM, so most work on muscle tissue involves thin section EM.
Production of thin sections involves the steps of fixation, embedding, sectioning followed by staining of the individual sections. Actin filaments are difficult to fix within muscle tissue because of their susceptibility to disruption by glutaraldehyde [ 36 ] and osmium [ 37 ], the conventional fixatives used for this purpose.
Osmium tetroxide is also a popular fixative for freeze substitution. To avoid this problem, Michael and Mary Reedy developed a fixative combining tannic acid and uranyl acetate, dubbed TAURAC, which avoided both glutaraldehyde and osmium treatments and proved to be compatible with freeze substitution [ 35 ]. Specimens prepared using TAURAC for the first time showed the helical structure of actin in X-ray fiber diffraction patterns of embedded muscle.
The development of a freeze substitution procedure using TAURAC opened the way to rapidly freezing active muscle for 3D visualization of myosin heads in action within the muscle lattice [ 35 ]. In the early s, the high degree of order within myac layer thin sections of the Lethocerus flight muscle suggested that 3D images could be obtained if the muscle lattice was treated as if it were a 2D protein array and images processed using methods developed for 2D crystals such as bacteriorhodopsin [ 38 ].
We refer to this approach as spatial averaging since the criterion used to decide if the repeating motifs unit cells can be averaged depends solely on their being periodically arranged within a lattice.
The missing data is commonly referred to as a missing wedge, cone or pyramid depending on the shape of the missing data. A single axis tilt series results in a missing wedge; a dual axis tilt series results in a missing pyramid of data. At the time, spatial averaging was the singular option for 3D imaging for this type of specimen. Crowther and Luther [ 23 ] developed a second spatial averaging option, the oblique section image reconstruction OSR , applicable to a specimen with a large unit cell for which sections could be cut substantially thinner than the unit cell dimensions.
Sections cut oblique to the muscle lattice are usually the default; cutting well-oriented myac layers e. The OSR was perfectly suited to 3D imaging of the Lethocerus flight muscle because it provided an average image of a complete unit cell, not just a single filament array, with no missing wedge. In some cases, the Fourier transform of the OSR could be compared with the X-ray diagram of the unfixed native muscle.
OSR was developed to its greatest extent using imaging of the Lethocerus flight muscle as the driving biological problem [ 11 , 26 , 28 , 39 , 40 ].
However, electron tomography ET with its ability to 3D image individual cross-bridges in situ and group classify similar structures regardless of position within the lattice ultimately replaced OSR. The application of ET to muscle thin sections solved most of the problems associated with intrinsic disorder in the filament lattice, as described below. In ET, the entire myac layer is reconstructed as a single volume.
Subvolume classification and averaging, which was a later development, were subsequently used to separate and group the different structures and arrangements prior to averaging [ 31 ]. Determination of the atomic structures of an actin monomer [ 41 ], the myosin head [ 27 ] and their combination into an atomic model of F-actin decorated with rigor myosin heads [ 42 ] initiated the process of atomic modeling of various states of the cross-bridge cycle within 3D images of muscle [ 32 , 43 ].
Atomic model building proved to be key for understanding the structure of myosin cross-bridges in situ. Although many crystal structures of myosin S1 or motor domains MD have been produced over the years, only one actin-bound state is characterized at high resolution, nucleotide-free acto-S1. Weak-binding actomyosin states are literally uncharacterized structurally. Weak actomyosin interactions occur frequently in muscle either with nucleotide analogs or as kinetic steps in active contractions.
The necessity of these modifications is, in fact, one of the more significant observations to emerge from 3D imaging actomyosin in situ. The Lethocerus flight muscle lattice, while well-ordered for a muscle, has certain kinds of intrinsic disorder that blur fine details in spatially averaged reconstructions.
One important disorder is the mismatch between helical spacings of the thick and thin filaments. The thick filament axial repeat is nm. The smallest common period is nm, of which there are about four full repeats in a half sarcomere, but four repeats is too few to be useful for averaging purposes.
The half pitch of the thin filament is Using the axial repeat of nm enabled the myosin and actin lattices to be reconstructed in the same operation. The azimuthal disorder generally prevents any actin subunits from being resolved in a spatially averaged reconstruction because two orientations offset axially by the actin subunit spacing of 2. Despite this, an important aspect of 3D images obtained by spatial averages was the ability to validate the observations using EMs of myac and actin layers.
The earliest 3D reconstruction work on muscle was confined to the rigor state. EM of thin sections had established the double chevron motif of the rigor Lethocerus flight muscle [ 1 , 5 ]. The double chevron Figure 1 C consisted of a large, angled cross-bridge pair towards the M-line, the lead chevron, separated from a smaller cross-bridge pair towards the Z-disk, the rear chevron, by a gap, the intra-doublet gap, that likely corresponded to a single actin subunit on each long pitch actin strand.
Of the actin subunits in each crossover, eight constituted the target zone of rigor muscle, four symmetrically placed on each long pitch strand. Later work as described below, showed that the two-actin subunits of the intra-doublet gap were rarely labeled by myosin heads regardless of the presence or absence of bound nucleotide. The eight-actin subunit target zone was later separated into a pair of lead- and a pair of rear-bridge target zones. Conversely, myosin head labeling of lead-bridge target zones retained some degree of order, but with greater heterogeneity.
Actin subunits were not resolved due to the thin filament intrinsic disorder, nor was the troponin complex identified. The troponin position was later identified by gold-Fab labeling which placed it near or at the rear chevron position [ 44 ]. The additional data from thick sections had the effect of better defining the size and shape of the cross-bridges and removing an apparent separation between the two long pitch strands of the thin filament.
The lead bridge took on a triangular shape with one edge of the triangle positioned on the thin filament; the second edge on the M-line side gave the lead bridge a strongly tilted appearance.
The third edge on the Z-disk side was oriented near perpendicular to the filament axis and one vertex of the triangle, the head-rod junction, was positioned on the thick filament surface e.
Although the two heads of the lead cross-bridge are not resolved, the shape is suggestive of one head, the leading, or M-line side head having a lever arm tilted at the classic rigor angle, and the trailing, or Z-line head, being less tilted toward the rigor and more perpendicular to the filament axis.
The region in the myac layer of A and the region of the actin layer of B contained in the transverse view in C are marked by the Wedgewood blue and teal green backgrounds respectively. L, Lead bridge; R, rear bridge; T, troponin. Triangles in B highlight the triangular shape of rigor lead bridges. Lead bridges of rigor and AMPPNP do not coincide exactly in shape so that in the superimposed region in the center, extra mass in rigor shows as copper color.
The troponin density is more pronounced in rigor, possibly because the thin filaments are better ordered. In B the cross-bridge pair that produces the two arms of the unflared-X are labeled 1 and 2. These same cross-bridges are similarly labeled in C. In C the rigor rear bridge is located at the far surface of the flared-X layer denoted by the arrow. The extra density provided by the rigor rear bridge results in the close proximity of the vertex arms in projections of flared-X.
Reprinted from [ 28 ] with permission from Elsevier. The OSR recapitulated these findings using a reconstruction procedure that suffers no missing pyramid such as occurs in a dual-axis tilt series, while reconstructing the protein density within a complete unit cell [ 40 ].
The OSR is also a spatial averaging technique so actin subunits could not be resolved. The resolution remained limited along the filament axis, but the advantages of the OSR outweighed this problem. Generally, the results of these various reconstructions on the rigor muscle indicated that the lead bridge consisted of two heads of a single myosin molecule diverging from a common origin at the head-rod junction and terminating on adjacent actin subunits on the thin filament.
The rear bridge was probably a single myosin head. Some rear bridge targets remained unlabeled by myosin heads. Myosin heads not bound to actin were disordered. Thus, myosin binding to lead-bridge actin targets appeared favored even in this early work and remained so in subsequent studies. Muscle states produced using non-hydrolysable ATP analogs AMPPNP , referred to as static states as opposed to dynamic states produced with ATP, were a research emphasis in the s—s before the development of rapid freezing techniques that could trap active cross-bridges.
Besides the rigor flight muscle, the only static states that had been characterized mechanically, by X-ray fiber diffraction and biochemically were those produced by the non-hydrolysable analogue AMPPNP with or without ethylene glycol [ 50 , 51 , 52 ]. The addition of either causes a drop in tension a measure of strong actin binding by myosin with little change in stiffness a measure of any actin binding, weak or strong, by myosin but together their effect is enhanced [ 51 ].
Thus, the calcium dependent status of the thin filament as well as temperature can influence this process. Because it changed the structure of rigor flight muscle myofibrils, as observed by both X-ray diffraction as well as EM, AMPPNP was a favorite subject for finding intermediate cross-bridge structures that might be related to changes occurring during muscle contraction.
However, the interpretation of AMPPNP effects were generally controversial regarding the binding affinity of attached and detached myosin heads which can be variable between myosin IIs from different species see [ 52 ] for a discussion of how these issues affect flight muscle. The X-ray diffraction indicated a state different from rigor or relaxed, but the EM of fixed, embedded, sectioned and stained fibers generally looked relaxed [ 55 ].
The relaxed appearance was shown to be a preparation artifact that could be avoided by careful monitoring by the X-ray fiber diffraction for changes induced by fixation [ 56 , 57 ]. The ratio I When mechanically monitored for changes in tension and stiffness, the aqueous AMPPNP addition reduced tension while retaining high stiffness [ 52 ].
The mechanical effects were not easily explained. Original images as well as optical filtered images of rigor and AMPPNP myac layers showed frequent cross-bridge binding in both states at a point midway between successive dark beads on the thin filament, the Tn complex.
The point between troponins where rigor lead bridges bind later proved to be the location of strong binding in active contraction [ 34 , 58 ]. X-ray studies on the Lethocerus flight muscle fibers in the active contraction show an I Micrographs of thin sections through these static states complement the 3D imaging. Two of these bridges were lead bridges and two were rear bridges of the double chevron [ 45 ]. Atomic model building of rigor lead and rear bridges in several studies indicated that the lever arm of the myosin head must be bent azimuthally to form the flared X, more so for the rear bridge than the lead bridge, suggesting considerable azimuthal flexibility in the myosin lever arm [ 32 , 43 ].
This return toward the 4-fold origins of myosin heads from the 2-fold origins in the rigor flared X would now be interpreted as a change in the lever arm azimuth possibly with changes in the position of the MD on the actin, described in more detail below.
Original images as well as the 3D reconstructions described below show that rigor-like rear bridges disappear when nucleotide is added and are replaced by much less regular myosin head attachments in the Tn region. Axial changes in the lever arm appear as an enhanced Domain 1 contains the actin-binding interface and Domain 2, which is close to the myosin surface, reflects the myosin head origin.
Domain 1, which we would now interpret as the MD, reports nucleotide-induced changes in its position on actin. Domain 2, which we would now interpret as the myosin lever arm, would reflect nucleotide-induced changes in the apparent origin at the thick filament surface.
The binding of nucleotide to actin-attached heads changes Domain 2 sufficiently to enable it to follow the helical origin of myosin heads, possibly in concert with a change in the position of Domain 1 on actin. The interpretation was compatible with a hypothesis that the myosin head consisted of two domains, one that bound actin tightly and was consequently immobilized, with another domain moving to produce muscle shortening [ 60 ].
The first myosin head crystal structure would not be solved for another six years [ 27 ] providing details of these two domains. The The strong density repeating on a The density corresponding to rear bridges of rigor is weak or absent in averaged images and best seen in original images where their variability was retained. The unflared X dominates images of 15 nm transverse sections. The OSR was done using a unique application of the technique, which utilized both transverse and longitudinal oblique sections to determine amplitudes and phases of the sampled Fourier transform [ 61 ] and is the most heavily averaged reconstruction done on the Lethocerus flight muscle.
The reconstruction showed little if any residual density at the position of the rigor rear bridge Figure 3 A,B and a lead cross-bridge that was less azimuthally bent than was typical of rigor Figure 3 C.
The lead bridge appearance agreed with original images as well as a tomographic reconstruction described below. However, the axial angle was less altered from the rigor angle than expected. Comparison of the structure factors Fourier coefficient amplitudes determined by the OSR with those determined by the X-ray fiber diffraction of the native tissue was one of the advantages of this reconstruction technique. Precise agreement between the same structure factor was unlikely due to the extreme difference between the specimen preparations, in one case fully hydrated and in the other embedded, sectioned and stained.
However, ratios of the same structure factors within the different preparations were informative. The comparison showed the underweight of the The retention of lead-type cross-bridges foretold the limitation of strong-binding cross-bridges in the actively contracting muscle to the rigor lead-bridge location. The lack of cross-bridges at the rigor rear bridge targets in AMPPNP could explain the large drop in tension, but not the retention of stiffness unless those heads detached by AMPPNP could reattach weakly to the thin filament with less ordering.
Atomic models built into 3D images of rigor rear cross-bridges indicated that the attached myosin heads would be highly strained compared with those of lead bridges [ 43 , 61 ] suggesting greater susceptibility to detachment by added nucleotide. Imaging of the Lethocerus flight muscle using averaged reconstruction methods had major limitations, which were always apparent, but not treatable at the time.
As long as a structure was regularly arranged, it would appear in the reconstruction, if not, it was either blurred or averaged out and not visible.
The intrinsic disorders prevented actin subunits and individual heads of 2-headed myosin attachments to actin from being resolved.
Although the OSR had some compensating advantages, e. ET came into its own as a structural technique when computer-controlled microscopes could collect tilt series automatically [ 29 ]. However, the earliest tomogram of the Lethocerus flight muscle [ 30 ] was obtained from the same dual axis tilt series used for spatial averaging [ 22 ] and thus had been manually recorded. The tilt series images were aligned using what is now referred to as marker free alignment, which was unique at the time and later developed into the general alignment tool contained in the program PROTOMO [ 64 , 65 ].
Interpretation required the independent development of subvolume alignment and classification to improve the signal-to-noise ratio and make these features interpretable in some detail [ 31 , 66 ]. Details on methods used for ET and subvolume averaging of the Lethocerus muscle can be obtained from several publications [ 35 , 67 , 68 , 69 ].
The first atomic model of actomyosin [ 42 ], which was obtained by a combination of an actomyosin cryoEM reconstruction with the atomic model of F-actin and the crystal structure of the myosin head, facilitated more detailed interpretation of the tomographic reconstructions.
An early development of quantitative atomic model refinement at low resolution was tested using subvolume averages derived from tomograms of rigor muscle [ 32 , 43 ]. In nearly every instance where a cross-bridge could be interpreted as binding actin strongly in situ, this model and more recent versions, required modification of the lever arm to facilitate a fit. Those modifications were greatest for the rigor rear bridge. Several modifications of the rigor Lethocerus flight muscle were imaged by ET including the rigor muscle swollen in low ionic strength buffer, which revealed the length of the proximal S2 domain that functions as a tether for the myosin heads [ 70 ].
Another study applied a stretch to the rigor muscle fibers followed by rapid freezing-freeze substitution to reveal elastic distortions in the myosin lever arm [ 33 ]. The tomograms of the rigor muscle recapitulated interpretations made earlier based on spatial averages Figure 4 A—C. Lead chevrons appeared in every Rear bridges were less regular and generally smaller when they were visible with their lever arms at variable angles.
When averaged, the rear bridges became a diffuse density on the Z-ward side of the lead chevron that in isodensity display appeared smaller and less angled than lead bridges. Multivariate data analysis of the repeating motifs, a process that can identify similar structures within a variable ensemble, had not been developed at this time. Instead, a novel averaging method dubbed column averaging was used.
In column averaging successive nm periods are averaged, but only along the axial direction; there is no lateral averaging between filaments. Column averages had less signal-to-noise improvement than spatial averages, but retained more of the cross-bridge variability.
The orientation has the Z-disc on the bottom. In B , E , H , the column-averaged filament is shown in red to the right of the unaveraged filaments gold on the left.
Target zone cross-bridges are colored red for ease of identification. The comparison clarifies the main differences among the three states; A — C the rigor state shows well-ordered, double chevrons consisting of lead- and rear-bridge pairs every There is no Weak White boxes outline mask motifs; G — I the glycol-PNP state shows single-headed attached cross-bridges every A mask motif is outlined by the white box.
The myosin filament surface reveals a strong Originally published in Journal of Cell Biology. In column averages, they appeared as a repeating pattern of apparently 2-headed and 1-headed lead bridges.
Rear type bridges disappeared even in the column averages, but were visible in the raw tomogram. Thus, the tension and stiffness effects of aqueous AMPPNP could be explained by the detachment of strong-binding rigor rear bridges that reattach weakly to the thin filament with disorder.
In rigor myac layers, paired rear bridges always bound the thin filament symmetrically about the filament axis and approached the thin filament from symmetric azimuths on the thick filament. In AMPPNP, this was not true as myosin heads contacting the rear bridge sites on the thin filament approached from nearly any direction on the thick filament, either front or back side. Their disordered appearance was consistent with nonspecific, weak attachments.
The availability of a rigor actomyosin atomic model facilitated a more detailed interpretation of the rigor tomograms. When placed in the reconstruction, MDs of the two actin-attached heads fit within the reconstruction envelope, but the lever arms of both heads would have to be adjusted azimuthally and axially if the two heads were to come together at a common origin, the head-rod junction [ 32 , 62 ], and still fit in the density. The fittings of the rigor rear bridge based on nucleotide-free acto-S1 demonstrate that in situ large lever arm deformations in the myosin head are possible when attaching actin in a state of high affinity.
Thus, the tomograms provided a direct observation consistent with the proposal that strain-enhanced detachment of cross-bridges explained the effects of AMPPNP in vertebrate preparations [ 71 , 72 ]; in flight muscle rear bridges were the most strained. As in the rigor tomogram, there was no clear indication where heads not attached to the thin filament were located.
Although in AMPPNP rigor rear bridges detached, they were reattaching to actin in ways not previously seen, but clearly different from the very specific attachment to actin found for strong binding myosin heads in rigor [ 62 ].
Subsequent information as described below indicates that M-ward bridges of mask motifs in AMPPNP must be very different from similar structures seen in the active muscle.
Adding ethylene glycol to the AMPPNP solution drove the muscle from rigor even more towards a relaxed appearance, while retaining stiffness nearly equal to that of rigor [ 50 ]. Tomograms of this glycol-stiff state in flight muscle fibers showed a significant structural change in the actin-bound heads Figure 4 G—I.
The actin azimuths for target zone cross-bridges in rigor and AMPPNP had a narrow distribution, the distribution of glycol stiff target zone cross-bridges was much broader, though still centered at the same average azimuth. Despite the large change in their structure, glycol-stiff, lead-bridge attachments retained the azimuthally symmetrical attachment to the actin targets characteristic of some specificity in their interaction with actin.
Mask motifs could still be observed in the glycol-stiff state Figure 4 H. Outside of the target zone, azimuthally, cross-bridges attached to just about every accessible surface of the thin filament. Thick filaments were marked by a strong The results from ET studies of rigor and these static states suggested that, as the affinity of the myosin head attaching to actin increased, the MD could position itself on actin independent of the lever arm, which must accommodate the thick filament origin as well as changes in the MD on actin [ 63 ].
This interpretation differed from other models that placed the MD on actin in all strong binding states in a single, stereospecific orientation with the lever arm moving only axially to produce filament sliding [ 42 ].
It also differed from models in which the entire myosin head rotated on actin during force production [ 2 , 73 ]. Several results published since the work described above for static, nucleotide-bound states of the Lethocerus flight muscle appeared to bear on their interpretation. These results include X-ray crystal structures of myosin II intermediate states obtained from MD constructs of Dictyostelium discoideum , high-resolution structures of actomyosin by cryoEM, and the structure of relaxed striated muscle thick filaments from several species.
The crystal structure of the D. Numerous crystal structures of complete myosin heads obtained from molluscan sources have similar features [ 78 , 79 , 80 , 81 ]. How do these results impact the tomographic reconstructions of the Lethocerus flight muscle in the presence of AMPPNP with or without ethylene glycol?
The free-head backbone attachment in the relaxed Lethocerus thick filaments is primarily through the RLC [ 6 ]; the position of the MD may be less important for this backbone attachment unless it clashes. Molluscan post-rigor myosin head structures, when aligned using the RLC and its bound heavy chain segment to the interacting heads motif of relaxed Lethocerus thick filaments, fit with no or only minor clashes between the MD and the thick filament backbone Figure 5.
The pyrophosphate addition to rigor the Lethocerus flight muscle also produces a structure similar to that of AMPPNP plus ethylene glycol with an enhanced Fitting of a post rigor myosin head conformation into the relaxed Lethocerus thick filament.
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Windows 10 1703 download iso itarget reviews google.Conclusion:
Rigor crossbridge structure in tilted single filament layers and flared-X formations from insect flight muscle. Skeletal muscle resists stretch by rapid binding of the second motor domain of myosin to actin. The file can be compressed but that also takes free disk space during compression. Dual axis tilt series tomogram of rigor Lethocerus flight muscle. Update the GPO that is deploying certificates by importing the required certificates,. Generally, the results of these various reconstructions on the rigor muscle indicated that the lead bridge consisted of two heads of a single myosin molecule diverging from a common origin at the head-rod junction and terminating on adjacent actin subunits on the thin filament. Developments in ET produced methods to align and classify the heterogeneous individual motifs ❿
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