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We are reliable and trusted among all our clients and thus you can entrust your academic work on us. C Increased apoptosis of Rictor -deficient B cells. D Competitive homeostasis of mature B cells. Mice were then treated with tamoxifen and were analyzed by flow cytometry 4 weeks after the adoptive transfer.
E Homeostatic proliferation of Rictor-deficient B cells. Shown is 1 result representative of those in 3 independent experiments, each consisting of 4 mice of each genotype.
Consistent with impaired Akt function at a nuclear target after Rictor depletion, much less phosphorylation of FoxO1 was observed Figure 5A. Shown are signal images for the indicated antibody in 1 of 3 independent experiments with comparable results. To explore the signaling defects in relation to B-cell survival, we analyzed expression of proapoptotic and antiapoptotic genes in B cells lacking Rictor.
These data indicate that Rictor assists in setting an appropriate balance of prosurvival and proapoptotic gene expression in B cells. Rictor regulates induction of prosurvival genes and suppression of proapoptotic genes. A Relative expression of Bcl2 family genes ex vivo.
Mature AA4. Antibody responses depend on the clonal expansion of antigen-reactive B cells, germinal center formation, and the differentiation and maintenance of plasma cells. This impact was most striking for high-affinity antibodies of class-switched isotypes. Moreover, germinal centers in the spleens of immunized Rictor-deficient mice were sparse and smaller supplemental Figure 4C. Comparison of these defects to our previous findings with depletion of Rictor in mature T cells 26 and the numbers of FO B-phenotype cells suggested that Rictor also acts within B cells in antibody responses.
High-affinity and class-switched antibodies are promoted by the germinal center reaction. Despite suitable T-cell help, frequencies of germinal center-phenotype B cells were reduced when Rictor-depleted B cells were used Figure 7D. Thus, we conclude that Rictor within B lineage cells promoted population expansion and the antibody response.
Overall, we propose that Rictor promotes the development of the preimmune B-cell populations followed by survival during and after clonal expansion. B-cell—intrinsic role of Rictor in antibody responses. Sera of these recipient mice were analyzed 3 weeks after immunization with NP-conjugated ovalbumin. Our data show that development, homeostasis, and immune function of mature B cells depend on Rictor, a protein central to mTORC2.
These defects, together with lower cell-cycling efficiency, suggest that B-cell—intrinsic contributions of mTORC2 to survival signaling and clonal expansion are major elements in the defective antibody responses. Intriguingly, TLR4-driven B-cell proliferation and in vitro antibody production exhibited less impact of Rictor deficiency Figure 4F ; supplemental Figure 6 , suggesting that antigen-driven, BCR-initiated signals may depend more stringently on Rictor than TLR-initiated processes.
It is revealing to integrate these findings with other aspects of PI3K signaling. Strikingly, the impact of Rictor on maturation appeared nearly as substantial as that of Akt 1, 2. Akt is catalytically active after activation loop phosphorylation by PDK1, but its activity and the persistence of Tphosphorylation are increased by phosphorylation of the conserved HM S of Akt.
The Akt3 isoform may be sufficiently functional in B lineage cells that the Akt phenotype was only partially penetrant, or the biochemical dose-response curve for Akt may require a level of catalytic activity not achieved without HM phosphorylation.
An intriguing possibility is that phosphorylation and structural rearrangement of the Akt HM promote functional targeting of biologically important substrates. FoxO1 deletion early in B lineage development caused blocks at pro- and pre-B—cell stages attributed to a failure to express RAG and the IL-7 receptor.
Recent findings with conditional Pten deletion show that the PI3K pathway affects the fetal blood progenitors very differently from stem cells of the adult marrow. Another striking phenotype observed in Rictor-deficient mice was a defect of MZ B cells, which were much more sensitive to acute deletion of Rictor than FO B cells.
Although PI3K is activated in a myriad of ways, it is intriguing that Notch 2 signaling both activates this lipid kinase and promotes MZ B-cell development, 50 because mTORC2 relays Notch signals in pre-T—cell differentiation and expansion. The online version of this article contains a data supplement. The publication costs of this article were defrayed in part by page charge payment. Contribution: K. The current address for K. Conflict-of-interest disclosure: The authors declare no competing financial interests.
Prepublished online Aug Thomas , 4 Robert C. Rickert , 2 and Mark Boothby 1, 4. James W. Robert C. Author information Article notes Copyright and License information Disclaimer. Corresponding author.
Received Jan 7; Accepted Aug This article has been cited by other articles in PMC. Rictor regulates survival of B cells and their balance of proapoptotic vs antiapoptotic gene expression. Introduction Humoral immunity relies on suitable pools of mature B-cell subsets, and their capacity for clonal expansion and differentiation into antibody-secreting cells.
Methods Detailed Methods are outlined in the supplemental Materials. Immunohistochemistry, B-cell isolation, and flow cytometry Immunohistochemistry was performed on frozen sections of spleens as described.
Western blotting; measurements of RNA Cell lysates were analyzed by immunoblotting. Open in a separate window. Figure 1. Figure 2. Figure 3. Figure 4. Figure 5. Figure 6. B-cell—intrinsic Rictor contribution to antibody responses Antibody responses depend on the clonal expansion of antigen-reactive B cells, germinal center formation, and the differentiation and maintenance of plasma cells.
Figure 7. Footnotes The online version of this article contains a data supplement. Authorship Contribution: K. References 1. B cell signaling and fate decision. Annu Rev Immunol. Schlissel MS. Regulating antigen-receptor gene assembly. Nat Rev Immunol. In vivo ablation of surface immunoglobulin on mature B cells by inducible gene targeting results in rapid cell death.
Clonal anergy: persistence in tolerant mice of antigen-binding B lymphocytes incapable of responding to antigen or mitogen. Shlomchik MJ, Weisel F.
Germinal center selection and the development of memory B and plasma cells. Immunol Rev. BAFF mediates survival of peripheral immature B lymphocytes. J Exp Med. BAFF binds to the tumor necrosis factor receptor-like molecule B cell maturation antigen and is important for maintaining the peripheral B cell population.
Maintenance of serum antibody levels. Impaired B and T cell antigen receptor signaling in pdelta PI 3-kinase mutant mice. Impaired B cell development and proliferation in absence of phosphoinositide 3-kinase p85alpha.
Fruman DA, Bismuth G. Fine tuning the immune response with PI3K. Vanhaesebroeck B, Alessi DR. Biochem J.
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The use of the cloud metaphor is credited to General Magic communications employee David Hoffman , based on long-standing use in networking and telecom.
In July , Amazon created subsidiary Amazon Web Services , with the goal to "enable developers to build innovative and entrepreneurial applications on their own. The goal was to eliminate the need for some administrative tasks typical of an IaaS model, while creating a platform where users could easily deploy such applications and scale them to demand. By mid, Gartner saw an opportunity for cloud computing "to shape the relationship among consumers of IT services, those who use IT services and those who sell them" [25] and observed that "organizations are switching from company-owned hardware and software assets to per-use service-based models" so that the "projected shift to computing In , the U.
National Science Foundation began the Cluster Exploratory program to fund academic research using Google - IBM cluster technology to analyze massive amounts of data.
The OpenStack project intended to help organizations offering cloud-computing services running on standard hardware. As an open-source offering and along with other open-source solutions such as CloudStack, Ganeti, and OpenNebula, it has attracted attention by several key communities. Several studies aim at comparing these open source offerings based on a set of criteria. On June 7, , Oracle announced the Oracle Cloud.
The goal of cloud computing is to allow users to take benefit from all of these technologies, without the need for deep knowledge about or expertise with each one of them. The cloud aims to cut costs and helps the users focus on their core business instead of being impeded by IT obstacles. Virtualization software separates a physical computing device into one or more "virtual" devices, each of which can be easily used and managed to perform computing tasks.
With operating system—level virtualization essentially creating a scalable system of multiple independent computing devices, idle computing resources can be allocated and used more efficiently. Virtualization provides the agility required to speed up IT operations and reduces cost by increasing infrastructure utilization. Autonomic computing automates the process through which the user can provision resources on-demand.
By minimizing user involvement, automation speeds up the process, reduces labor costs and reduces the possibility of human errors. Cloud computing uses concepts from utility computing to provide metrics for the services used. Cloud computing attempts to address QoS quality of service and reliability problems of other grid computing models. The National Institute of Standards and Technology 's definition of cloud computing identifies "five essential characteristics":.
On-demand self-service. A consumer can unilaterally provision computing capabilities, such as server time and network storage, as needed automatically without requiring human interaction with each service provider. Broad network access. Capabilities are available over the network and accessed through standard mechanisms that promote use by heterogeneous thin or thick client platforms e. Resource pooling. The provider's computing resources are pooled to serve multiple consumers using a multi-tenant model, with different physical and virtual resources dynamically assigned and reassigned according to consumer demand.
Rapid elasticity. Capabilities can be elastically provisioned and released, in some cases automatically, to scale rapidly outward and inward commensurate with demand. To the consumer, the capabilities available for provisioning often appear unlimited and can be appropriated in any quantity at any time.
Measured service. Cloud systems automatically control and optimize resource use by leveraging a metering capability at some level of abstraction appropriate to the type of service e. Resource usage can be monitored, controlled, and reported, providing transparency for both the provider and consumer of the utilized service. For example, one can provide SaaS implemented on physical machines bare metal , without using underlying PaaS or IaaS layers, and conversely one can run a program on IaaS and access it directly, without wrapping it as SaaS.
A hypervisor runs the virtual machines as guests. Pools of hypervisors within the cloud operational system can support large numbers of virtual machines and the ability to scale services up and down according to customers' varying requirements. Linux containers run in isolated partitions of a single Linux kernel running directly on the physical hardware. Linux cgroups and namespaces are the underlying Linux kernel technologies used to isolate, secure and manage the containers.
Containerisation offers higher performance than virtualization because there is no hypervisor overhead. IaaS clouds often offer additional resources such as a virtual-machine disk-image library, raw block storage , file or object storage , firewalls, load balancers, IP addresses , virtual local area networks VLANs , and software bundles.
The NIST 's definition of cloud computing describes IaaS as "where the consumer is able to deploy and run arbitrary software, which can include operating systems and applications.
The consumer does not manage or control the underlying cloud infrastructure but has control over operating systems, storage, and deployed applications; and possibly limited control of select networking components e.
IaaS-cloud providers supply these resources on-demand from their large pools of equipment installed in data centers. For wide-area connectivity, customers can use either the Internet or carrier clouds dedicated virtual private networks.
To deploy their applications, cloud users install operating-system images and their application software on the cloud infrastructure. In this model, the cloud user patches and maintains the operating systems and the application software. Cloud providers typically bill IaaS services on a utility computing basis: cost reflects the number of resources allocated and consumed.
The capability provided to the consumer is to deploy onto the cloud infrastructure consumer-created or acquired applications created using programming languages, libraries, services, and tools supported by the provider. The consumer does not manage or control the underlying cloud infrastructure including network, servers, operating systems, or storage, but has control over the deployed applications and possibly configuration settings for the application-hosting environment.
PaaS vendors offer a development environment to application developers. The provider typically develops toolkit and standards for development and channels for distribution and payment.
In the PaaS models, cloud providers deliver a computing platform , typically including an operating system, programming-language execution environment, database, and the web server. Application developers develop and run their software on a cloud platform instead of directly buying and managing the underlying hardware and software layers. With some PaaS, the underlying computer and storage resources scale automatically to match application demand so that the cloud user does not have to allocate resources manually.
Some integration and data management providers also use specialized applications of PaaS as delivery models for data. The capability provided to the consumer is to use the provider's applications running on a cloud infrastructure. The applications are accessible from various client devices through either a thin client interface, such as a web browser e. The consumer does not manage or control the underlying cloud infrastructure including network, servers, operating systems, storage, or even individual application capabilities, with the possible exception of limited user-specific application configuration settings.
In the software as a service SaaS model, users gain access to application software and databases. Cloud providers manage the infrastructure and platforms that run the applications. SaaS is sometimes referred to as "on-demand software" and is usually priced on a pay-per-use basis or using a subscription fee. Cloud users do not manage the cloud infrastructure and platform where the application runs. This eliminates the need to install and run the application on the cloud user's own computers, which simplifies maintenance and support.
Cloud applications differ from other applications in their scalability—which can be achieved by cloning tasks onto multiple virtual machines at run-time to meet changing work demand.
This process is transparent to the cloud user, who sees only a single access-point. To accommodate a large number of cloud users, cloud applications can be multitenant , meaning that any machine may serve more than one cloud-user organization. The pricing model for SaaS applications is typically a monthly or yearly flat fee per user, [79] so prices become scalable and adjustable if users are added or removed at any point.
It may also be free. In addition, with applications hosted centrally, updates can be released without the need for users to install new software.
One drawback of SaaS comes with storing the users' data on the cloud provider's server. As a result, [ citation needed ] there could be unauthorized access to the data. SaaS applications may be integrated with cloud storage or File hosting services , which is the case with Google Docs being integrated with Google Drive and Word Online being integrated with Onedrive. In the mobile "backend" as a service m model, also known as backend as a service BaaS , web app and mobile app developers are provided with a way to link their applications to cloud storage and cloud computing services with application programming interfaces APIs exposed to their applications and custom software development kits SDKs.
Services include user management, push notifications , integration with social networking services [82] and more. This is a relatively recent model in cloud computing, [83] with most BaaS startups dating from or later [84] [85] [86] but trends indicate that these services are gaining significant mainstream traction with enterprise consumers.
Serverless computing is a cloud computing code execution model in which the cloud provider fully manages starting and stopping virtual machines as necessary to serve requests, and requests are billed by an abstract measure of the resources required to satisfy the request, rather than per virtual machine, per hour. Function as a service FaaS is a service-hosted remote procedure call that leverages serverless computing to enable the deployment of individual functions in the cloud that run in response to events.
Private cloud is cloud infrastructure operated solely for a single organization, whether managed internally or by a third party, and hosted either internally or externally. It can improve business, but every step in the project raises security issues that must be addressed to prevent serious vulnerabilities. Self-run data centers [91] are generally capital intensive.
They have a significant physical footprint, requiring allocations of space, hardware, and environmental controls. These assets have to be refreshed periodically, resulting in additional capital expenditures. They have attracted criticism because users "still have to buy, build, and manage them" and thus do not benefit from less hands-on management, [92] essentially "[lacking] the economic model that makes cloud computing such an intriguing concept".
Cloud services are considered "public" when they are delivered over the public Internet, and they may be offered as a paid subscription, or free of charge. Most public-cloud providers offer direct-connection services that allow customers to securely link their legacy data centers to their cloud-resident applications. Hybrid cloud is a composition of a public cloud and a private environment, such as a private cloud or on-premises resources, [98] [99] that remain distinct entities but are bound together, offering the benefits of multiple deployment models.
It allows one to extend either the capacity or the capability of a cloud service, by aggregation, integration or customization with another cloud service.
Varied use cases for hybrid cloud composition exist. For example, an organization may store sensitive client data in house on a private cloud application, but interconnect that application to a business intelligence application provided on a public cloud as a software service.
Hybrid cloud adoption depends on a number of factors such as data security and compliance requirements, level of control needed over data, and the applications an organization uses.
Another example of hybrid cloud is one where IT organizations use public cloud computing resources to meet temporary capacity needs that can not be met by the private cloud. A primary advantage of cloud bursting and a hybrid cloud model is that an organization pays for extra compute resources only when they are needed.
A cross-platform hybrid cloud is usually powered by different CPU architectures, for example, x and ARM, underneath. Users can transparently deploy and scale applications without knowledge of the cloud's hardware diversity. Hybrid cloud infrastructure essentially serves to eliminate limitations inherent to the multi-access relay characteristics of private cloud networking. The advantages include enhanced runtime flexibility and adaptive memory processing unique to virtualized interface models.
Community cloud shares infrastructure between several organizations from a specific community with common concerns security, compliance, jurisdiction, etc. The costs are spread over fewer users than a public cloud but more than a private cloud , so only some of the cost savings potential of cloud computing are realized. A cloud computing platform can be assembled from a distributed set of machines in different locations, connected to a single network or hub service. It is possible to distinguish between two types of distributed clouds: public-resource computing and volunteer cloud.
Multicloud is the use of multiple cloud computing services in a single heterogeneous architecture to reduce reliance on single vendors, increase flexibility through choice, mitigate against disasters, etc. It differs from hybrid cloud in that it refers to multiple cloud services, rather than multiple deployment modes public, private, legacy.
Poly cloud refers to the use of multiple public clouds for the purpose of leveraging specific services that each provider offers. It differs from Multi cloud in that it is not designed to increase flexibility or mitigate against failures but is rather used to allow an organization to achieve more that could be done with a single provider.
The issues of transferring large amounts of data to the cloud as well as data security once the data is in the cloud initially hampered adoption of cloud for big data , but now that much data originates in the cloud and with the advent of bare-metal servers , the cloud has become [] a solution for use cases including business analytics and geospatial analysis. HPC cloud refers to the use of cloud computing services and infrastructure to execute high-performance computing HPC applications.
Cloud architecture , [] the systems architecture of the software systems involved in the delivery of cloud computing, typically involves multiple cloud components communicating with each other over a loose coupling mechanism such as a messaging queue. Elastic provision implies intelligence in the use of tight or loose coupling as applied to mechanisms such as these and others. Cloud engineering is the application of engineering disciplines of cloud computing. It brings a systematic approach to the high-level concerns of commercialization, standardization and governance in conceiving, developing, operating and maintaining cloud computing systems.
It is a multidisciplinary method encompassing contributions from diverse areas such as systems , software , web , performance , information technology engineering , security , platform , risk , and quality engineering. Cloud computing poses privacy concerns because the service provider can access the data that is in the cloud at any time.
It could accidentally or deliberately alter or delete information. That is permitted in their privacy policies, which users must agree to before they start using cloud services. Solutions to privacy include policy and legislation as well as end-users' choices for how data is stored. These systems distinguish between authorized and unauthorized users and determine the amount of data that is accessible to each entity.
Together, these form shared technology vulnerabilities. In a cloud provider platform being shared by different users, there may be a possibility that information belonging to different customers resides on the same data server.
Additionally, Eugene Schultz , chief technology officer at Emagined Security, said that hackers are spending substantial time and effort looking for ways to penetrate the cloud. Because data from hundreds or thousands of companies can be stored on large cloud servers, hackers can theoretically gain control of huge stores of information through a single attack—a process he called "hyperjacking".
Some examples of this include the Dropbox security breach, and iCloud leak. By having these passwords, they are able to read private data as well as have this data be indexed by search engines making the information public. There is the problem of legal ownership of the data If a user stores some data in the cloud, can the cloud provider profit from it? Many Terms of Service agreements are silent on the question of ownership. This delivers great incentive to public cloud computing service providers to prioritize building and maintaining strong management of secure services.
There is the risk that end users do not understand the issues involved when signing on to a cloud service persons sometimes don't read the many pages of the terms of service agreement, and just click "Accept" without reading. This is important now that cloud computing is common and required for some services to work, for example for an intelligent personal assistant Apple's Siri or Google Assistant.
Fundamentally, private cloud is seen as more secure with higher levels of control for the owner, however public cloud is seen to be more flexible and requires less time and money investment from the user.
According to Bruce Schneier , "The downside is that you will have limited customization options. Cloud computing is cheaper because of economics of scale , and—like any outsourced task—you tend to get what you want.
A restaurant with a limited menu is cheaper than a personal chef who can cook anything you want. Fewer options at a much cheaper price: it's a feature, not a bug.
Cloud providers often decide on the management policies, which moderates what the cloud users are able to do with their deployment. Privacy and confidentiality are big concerns in some activities.
For instance, sworn translators working under the stipulations of an NDA , might face problems regarding sensitive data that are not encrypted. Cloud computing has some limitations for smaller business operations, particularly regarding security and downtime. Technical outages are inevitable and occur sometimes when cloud service providers CSPs become overwhelmed in the process of serving their clients.
This may result in temporary business suspension. Since this technology's systems rely on the Internet, an individual cannot access their applications, server, or data from the cloud during an outage. Cloud computing is still a subject of research. Since the global pandemic of , cloud technology jumped ahead in popularity due to the level of security of data and the flexibility of working options for all employees, notably remote workers. The issue of carrying out investigations where the cloud storage devices cannot be physically accessed has generated a number of changes to the way that digital evidence is located and collected.
In some scenarios existing digital forensics tools can be employed to access cloud storage as networked drives although this is a slow process generating a large amount of internet traffic. An alternative approach is to deploy a tool that processes in the cloud itself. For organizations using Office with an 'E5' subscription, there is the option to use Microsoft's built-in e-discovery resources, although these do not provide all the functionality that is typically required for a forensic process.
From Wikipedia, the free encyclopedia. Form of shared Internet-based computing. Not to be confused with Cloud Computing horse. This article may be confusing or unclear to readers. A Excision of Rictor in B lymphoid cells. D Flow cytometry of spleen cells in the viable lymphoid gate or further gates as indicated above panels.
Vav- Cre also deletes the conditional Rictor allele outside of the B lineage. Marrow programmed to delete Rictor exhibited a competitive disadvantage in generation of mature splenic B cells compared with WT controls Figure 2A.
Despite a hematopoietic compartment generating normal lineages WT CD However, these decreases were modest compared with the impact on mature B cells. Collectively, these findings demonstrate a cell-autonomous role for Rictor in the establishment of mature B-cell populations. Defects in repopulation of Rictor-deficient B cells in a competitive fitness model. Ratio of CD Shown are representative fluorescence-activated cell sorter FACS profiles in the gates as indicated in the above panels right.
C Ratios of CD No defect was apparent 2 weeks after the first tamoxifen injection data not shown. Although Rictor deletion impairs thymic production of new T cells, 29 B-cell maintenance is T cell independent.
Together, these results indicate that Rictor mediates maintenance of mature B cells. Effect of acute Rictor deletion on B lymphoid cells.
C Representative FACS profiles with frequencies of the indicated subsets in the viable lymphoid gates of the spleen or further gates as indicated in the above panels. Thus, this protein assists survival signaling in B cells. Impaired survival, proliferation, and persistence of Rictor-deficient B cells. C Increased apoptosis of Rictor -deficient B cells. D Competitive homeostasis of mature B cells. Mice were then treated with tamoxifen and were analyzed by flow cytometry 4 weeks after the adoptive transfer.
E Homeostatic proliferation of Rictor-deficient B cells. Shown is 1 result representative of those in 3 independent experiments, each consisting of 4 mice of each genotype. Consistent with impaired Akt function at a nuclear target after Rictor depletion, much less phosphorylation of FoxO1 was observed Figure 5A.
Shown are signal images for the indicated antibody in 1 of 3 independent experiments with comparable results. To explore the signaling defects in relation to B-cell survival, we analyzed expression of proapoptotic and antiapoptotic genes in B cells lacking Rictor. These data indicate that Rictor assists in setting an appropriate balance of prosurvival and proapoptotic gene expression in B cells. Rictor regulates induction of prosurvival genes and suppression of proapoptotic genes.
A Relative expression of Bcl2 family genes ex vivo. Mature AA4. Antibody responses depend on the clonal expansion of antigen-reactive B cells, germinal center formation, and the differentiation and maintenance of plasma cells.
This impact was most striking for high-affinity antibodies of class-switched isotypes. Moreover, germinal centers in the spleens of immunized Rictor-deficient mice were sparse and smaller supplemental Figure 4C.
Comparison of these defects to our previous findings with depletion of Rictor in mature T cells 26 and the numbers of FO B-phenotype cells suggested that Rictor also acts within B cells in antibody responses. High-affinity and class-switched antibodies are promoted by the germinal center reaction.
Despite suitable T-cell help, frequencies of germinal center-phenotype B cells were reduced when Rictor-depleted B cells were used Figure 7D. Thus, we conclude that Rictor within B lineage cells promoted population expansion and the antibody response. Overall, we propose that Rictor promotes the development of the preimmune B-cell populations followed by survival during and after clonal expansion. B-cell—intrinsic role of Rictor in antibody responses. Sera of these recipient mice were analyzed 3 weeks after immunization with NP-conjugated ovalbumin.
Our data show that development, homeostasis, and immune function of mature B cells depend on Rictor, a protein central to mTORC2. These defects, together with lower cell-cycling efficiency, suggest that B-cell—intrinsic contributions of mTORC2 to survival signaling and clonal expansion are major elements in the defective antibody responses.
Intriguingly, TLR4-driven B-cell proliferation and in vitro antibody production exhibited less impact of Rictor deficiency Figure 4F ; supplemental Figure 6 , suggesting that antigen-driven, BCR-initiated signals may depend more stringently on Rictor than TLR-initiated processes.
It is revealing to integrate these findings with other aspects of PI3K signaling. Strikingly, the impact of Rictor on maturation appeared nearly as substantial as that of Akt 1, 2. Akt is catalytically active after activation loop phosphorylation by PDK1, but its activity and the persistence of Tphosphorylation are increased by phosphorylation of the conserved HM S of Akt.
The Akt3 isoform may be sufficiently functional in B lineage cells that the Akt phenotype was only partially penetrant, or the biochemical dose-response curve for Akt may require a level of catalytic activity not achieved without HM phosphorylation.
An intriguing possibility is that phosphorylation and structural rearrangement of the Akt HM promote functional targeting of biologically important substrates. FoxO1 deletion early in B lineage development caused blocks at pro- and pre-B—cell stages attributed to a failure to express RAG and the IL-7 receptor.
Recent findings with conditional Pten deletion show that the PI3K pathway affects the fetal blood progenitors very differently from stem cells of the adult marrow.
Another striking phenotype observed in Rictor-deficient mice was a defect of MZ B cells, which were much more sensitive to acute deletion of Rictor than FO B cells. Although PI3K is activated in a myriad of ways, it is intriguing that Notch 2 signaling both activates this lipid kinase and promotes MZ B-cell development, 50 because mTORC2 relays Notch signals in pre-T—cell differentiation and expansion.
The online version of this article contains a data supplement. The publication costs of this article were defrayed in part by page charge payment. Contribution: K. The current address for K. Conflict-of-interest disclosure: The authors declare no competing financial interests. Prepublished online Aug Thomas , 4 Robert C.
Rickert , 2 and Mark Boothby 1, 4. James W. Robert C. Author information Article notes Copyright and License information Disclaimer. Corresponding author. Received Jan 7; Accepted Aug This article has been cited by other articles in PMC. Rictor regulates survival of B cells and their balance of proapoptotic vs antiapoptotic gene expression. Introduction Humoral immunity relies on suitable pools of mature B-cell subsets, and their capacity for clonal expansion and differentiation into antibody-secreting cells.
Methods Detailed Methods are outlined in the supplemental Materials. Immunohistochemistry, B-cell isolation, and flow cytometry Immunohistochemistry was performed on frozen sections of spleens as described.
Western blotting; measurements of RNA Cell lysates were analyzed by immunoblotting. Open in a separate window. Figure 1. Figure 2. Figure 3. Figure 4. Figure 5.
Figure 6. B-cell—intrinsic Rictor contribution to antibody responses Antibody responses depend on the clonal expansion of antigen-reactive B cells, germinal center formation, and the differentiation and maintenance of plasma cells.
Figure 7. Footnotes The online version of this article contains a data supplement. Authorship Contribution: K. References 1. B cell signaling and fate decision.
Annu Rev Immunol.
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