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|By Elizabeth White
Article Rating:
|September 2, 2014 10:45 PM EDT
Compuware Corporation (Nasdaq:CPWR) and Thoma Bravo jointly announced that Compuware has entered into a definitive agreement to be acquired by private equity investment firm Thoma Bravo, LLC, in a transaction valued at approximately $2.5 billion.
"Compuware is the clear established leader in the categories of application performance and mainframe productivity tools, and this transaction is the capstone to a series of transformative company initiatives to relentlessly drive value," said Bob Paul, Chief Executive Officer of Compuware. "We began with the IPO of Covisint, initiated a robust dividend, divested non-core operations, and aggressively reduced corporate expenses. Compuware is now best suited to focus on its core mainframe and APM businesses as a private-equity backed company, where we can continue to serve our customers in a competitive environment with greater flexibility to take a long-term approach."
"Organizations are increasingly relying on mission-critical technologies and applications to reach customers and grow their businesses, and Compuware's solutions, including Compuware APM, are the leading choice by many of the world's largest organizations for ensuring those applications perform seamlessly," said Orlando Bravo, a managing partner at Thoma Bravo. "Becoming a private company will enable this established market leader to leverage strategic product and other growth opportunities that will take Compuware to the next level."
Under the terms of the agreement, pending shareholder approval, Compuware shareholders will receive an aggregate value of approximately $10.92 per share, representing a premium of approximately 17 percent to the Company's stock price as of the close of trading on Friday, August 29, 2014. Thoma Bravo will pay a cash purchase price of $10.43 for each outstanding share of Compuware common stock, less the pro rata portion of the applicable corporate tax that will be owed in connection with the spin-off of Covisint, currently estimated at $0.18 per share based on the current market price of Covisint, for a net cash payment of approximately $10.25 per share. The parties have agreed that within 60 days following the date of the merger agreement, Compuware will effectuate the pro rata distribution to its shareholders of the remaining shares of Covisint owned by Compuware, resulting in a distribution of Covisint shares representing approximately $0.67 per share of Compuware common stock based on the closing price of Covisint on Friday, August 29. During the 60-day period, Compuware may seek a higher value alternative for its Covisint shares, in which case the proceeds, net of tax and certain charges, of such disposition will be paid to Compuware shareholders.
Gurminder S. Bedi, Independent Chairman of the Compuware Board, said, "This is the right transaction for Compuware at the right time, and reflects a thorough Board review of strategic alternatives and the work of a committee established earlier this year to focus on value-generating steps. This agreement provides shareholders with immediate and substantial cash value, a significant premium to our share price, and the ability to complete the Covisint spin-off to shareholders. Thoma Bravo is an ideal partner for Compuware, adding significant application software, services, and financial expertise."
"We have been incredibly impressed with the business that the Compuware management team has built, and look forward to working with them on this next stage of growth for the company," added Seth Boro, a managing partner at Thoma Bravo. "The APM and Mainframe Productivity Tools markets are exciting and ever-evolving industries, and we're confident that our partnership with Compuware will enhance its position as the market leader and fuel further innovation that will benefit customers."
The Compuware board of directors unanimously approved the agreement and recommends that Compuware's shareholders approve the transaction. The transaction, which is expected to close by early 2015, is subject to approval from Compuware's shareholders, regulatory approvals, and other customary closing conditions. The closing of the transaction is also subject to the completion of a disposition of Covisint.
Elliott Management, which owns approximately 9.5 percent of Compuware's common stock, has entered into an agreement with Thoma Bravo agreeing to vote its shares in favor of the transaction.
There is no financing condition associated with the proposed acquisition. Jefferies, Credit Suisse and Deutsche Bank have agreed to provide debt financing in connection with the transaction.
Compuware has agreed to immediately discontinue its quarterly cash dividend. At closing, Thoma Bravo will acquire 100% of Compuware's outstanding shares and Compuware will become a privately-held company.
Goldman, Sachs & Co. is serving as financial advisor, and Skadden, Arps, Slate, Meagher & Flom LLP is serving as legal counsel to Compuware. Kirkland & Ellis LLP is serving as legal counsel to Thoma Bravo.
Cloud Expo's giant Silicon View billboard which is viewed by more than 1.3 million motorists per week
The 15th International Cloud Expo has just expanded its conference program, to bring together Cloud Computing, APM, APIs, Security, Big Data, Internet of Things, DevOps and WebRTC at one location.
The show now has three tracks devoted exclusively to the IoT (with WebRTC present in one of the tracks), a full single track focusing on Big Data, and a two-track DevOps Summit, in addition to four tracks devoted exclusively to Cloud Computing in the enterprise.
Cloud Expo is the single show where delegates and technology vendors can meet to experience and discuss the entire world of the cloud.
With Cloud Computing driving a higher percentage of enterprise IT budgets every year, it becomes increasingly important to learn about the latest technology developments and solutions.
Attend Cloud Expo Nov 4-6, at the Santa Clara Convention Center, CA. Craft your own custom experience. Learn the latest from the world's best technologists. Find the vendors you want and put them to the test.
15th Cloud Expo Expanded Conference Program
Cloud Expo
Track 1: Enterprise Cloud Adoption
Track 2: APM, Cloud and Hot Topics
Track 3: Cloud APIs & Business
Track 4: Cloud Security - Mobility
Big Data Expo
Track 5: Big Data/Analytics
Track 6: Consumer IoT
Track 7: Enterprise IoT
Track 8: IoT Developer & WebRTC
DevOps Summit
Track 1: "Dev" Developer Focus
Track 2: "Ops" Operations Focus
More than 150 all-star faculty members will discuss the hottest technology topics in November. In addition to the expanded APM and API coverage, Cloud Expo Silicon Valley will offer a variety of Linux containers and Docker sessions.
DevOps Summit at Cloud Expo also doubled its content from June, in two full simultaneous tracks for three days.
Tracks and topics for DevOps Summit at Cloud Expo will include:
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- ROI and business value
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- Quality assurance
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- DevOps and ITIL, ISO, Six Sigma, COBIT
- Mobile DevOps
- DevOps for legacy systems
- Software-defined servers
- Network automation
- Server automation
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- DevOps anti-patterns
- Enterprise DevOps
- DevOps system architecture
- IT orchestration
- Containerization
- DevOps skills and training
- WebOps, CloudOps, ChatOps, NoOps
- Change management
Tracks and topics for Internet of @ThingsExpo will include:
Consumer IoT & Wearables
- Smart Appliances
- Smart Cars
- Smartphones 2.0
- Smart Travel
- Personal Fitness
- Health Care
- Personalized Marketing
- Customized Shopping
- Personal Finance
- The Digital Divide
- Mobile Cash & Markets
- Games & the IoT
- The Future of Education
- Virtual Reality
Enterprise IoT
- The Business Case for IoT
- Smart Grids
- Smart Cities
- Smart Transportation
- The Smart Home
- Authentication/Security
- Wiring the IoT
- The Internet of Everything
- Digital Transformation of Enterprise IT
- Agriculture
- Transportation
- Manufacturing
- Local & State Government
- Federal Government
IoT Developers
- Eclipse Foundation
- Cloud Foundry
- Linux Containers
- Node-Red
- Open Source Hardware
- AJAX and the IoT
- Leveraging SOA
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- Nest Labs
Sponsorship Opportunities
for Cloud Expo® 2014 Silicon Valley
Cloud Expo® 2014 Silicon Valley "show prospectus" has shipped. Sponsorship, exhibit, and keynote opportunities can be obtained from Carmen Gonzalez by email at events (at) sys-con.com, or by phone 201 802-3021.
Early Bird Registration Options
for Cloud Expo® 2014 Silicon Valley
Cloud Expo® delegates can pre-register for Cloud Expo® Silicon Valley with Early Bird savings here.
About SYS-CON Media & Events
SYS-CON Media (www.sys-con.com) has since 1994 been connecting technology companies and customers through a comprehensive content stream at www.sys-con.com - featuring over forty focused subject areas, from Cloud Computing to Web Security - interwoven with market-leading full-scale conferences produced by SYS-CON Events. The company's internationally recognized brands include among others Cloud Expo® (www.CloudComputingExpo.com), Big Data Expo (http://BigDataExpo.net), Virtualization Conference & Expo (www.VirtualizationConference.com), DevOps Summit (http://DevOpsSummit.sys-con.com), Internet of @ThingsExpo (http://www.ThingsExpo.com), SDDC Expo (http://SDDCExpo.sys-con.com), WebRTC Summit (http://WebRTCSummit.net), Government IT Conference & Expo (www.GovITExpo.com), Cloud Computing Bootcamp (www.CloudComputingBootcamp.com), and UlitzerLive! New Media Conference & Expo (http://events.sys-con.com).
Cloud Expo® is a registered trademark of Cloud Expo, Inc.
The buzz continues for cloud, data analytics and the Internet of Things (IoT) and their collective impact across all industries. But a new conversation is emerging - how do companies use industry disruption and technology enablers to lead in markets undergoing change, uncertainty and ambiguity? Organizations of all sizes need to evolve and transform, often under massive pressure, as industry lines blur and merge and traditional business models are assaulted and turned upside down. In this new data-driven world, marketplaces reign supreme while interoperability, APIs and applications deliver un...
Oct. 9, 2015 08:00 AM EDT Reads: 285
Too often with compelling new technologies market participants become overly enamored with that attractiveness of the technology and neglect underlying business drivers. This tendency, what some call the "newest shiny object syndrome," is understandable given that virtually all of us are heavily engaged in technology. But it is also mistaken. Without concrete business cases driving its deployment, IoT, like many other technologies before it, will fade into obscurity.
Oct. 9, 2015 08:00 AM EDT
Today air travel is a minefield of delays, hassles and customer disappointment. Airlines struggle to revitalize the experience. GE and M2Mi will demonstrate practical examples of how IoT solutions are helping airlines bring back personalization, reduce trip time and improve reliability. In their session at @ThingsExpo, Shyam Varan Nath, Principal Architect with GE, and Dr. Sarah Cooper, M2Mi's VP Business Development and Engineering, will explore the IoT cloud-based platform technologies driving this change including privacy controls, data transparency and integration of real time context w...
Oct. 9, 2015 07:30 AM EDT
"Matrix is an ambitious open standard and implementation that's set up to break down the fragmentation problems that exist in IP messaging and VoIP communication," explained John Woolf, Technical Evangelist at Matrix, in this SYS-CON.tv interview at @ThingsExpo, held Nov 4–6, 2014, at the Santa Clara Convention Center in Santa Clara, CA.
Oct. 9, 2015 07:00 AM EDT Reads: 5,880
WebRTC converts the entire network into a ubiquitous communications cloud thereby connecting anytime, anywhere through any point. In his session at WebRTC Summit,, Mark Castleman, EIR at Bell Labs and Head of Future X Labs, will discuss how the transformational nature of communications is achieved through the democratizing force of WebRTC. WebRTC is doing for voice what HTML did for web content.
Oct. 9, 2015 06:00 AM EDT Reads: 1,411
The IoT is upon us, but today's databases, built on 30-year-old math, require multiple platforms to create a single solution. Data demands of the IoT require Big Data systems that can handle ingest, transactions and analytics concurrently adapting to varied situations as they occur, with speed at scale. In his session at @ThingsExpo, Chad Jones, chief strategy officer at Deep Information Sciences, will look differently at IoT data so enterprises can fully leverage their IoT potential. He'll share tips on how to speed up business initiatives, harness Big Data and remain one step ahead by apply...
Oct. 9, 2015 05:15 AM EDT Reads: 511
Nowadays, a large number of sensors and devices are connected to the network. Leading-edge IoT technologies integrate various types of sensor data to create a new value for several business decision scenarios. The transparent cloud is a model of a new IoT emergence service platform. Many service providers store and access various types of sensor data in order to create and find out new business values by integrating such data.
Oct. 9, 2015 04:00 AM EDT Reads: 570
The broad selection of hardware, the rapid evolution of operating systems and the time-to-market for mobile apps has been so rapid that new challenges for developers and engineers arise every day. Security, testing, hosting, and other metrics have to be considered through the process. In his session at Big Data Expo, Walter Maguire, Chief Field Technologist, HP Big Data Group, at Hewlett-Packard, will discuss the challenges faced by developers and a composite Big Data applications builder, focusing on how to help solve the problems that developers are continuously battling.
Oct. 9, 2015 04:00 AM EDT Reads: 500
There are so many tools and techniques for data analytics that even for a data scientist the choices, possible systems, and even the types of data can be daunting. In his session at @ThingsExpo, Chris Harrold, Global CTO for Big Data Solutions for EMC Corporation, will show how to perform a simple, but meaningful analysis of social sentiment data using freely available tools that take only minutes to download and install. Participants will get the download information, scripts, and complete end-to-end walkthrough of the analysis from start to finish. Participants will also be given the pract...
Oct. 9, 2015 03:00 AM EDT Reads: 293
WebRTC services have already permeated corporate communications in the form of videoconferencing solutions. However, WebRTC has the potential of going beyond and catalyzing a new class of services providing more than calls with capabilities such as mass-scale real-time media broadcasting, enriched and augmented video, person-to-machine and machine-to-machine communications. In his session at @ThingsExpo, Luis Lopez, CEO of Kurento, will introduce the technologies required for implementing these ideas and some early experiments performed in the Kurento open source software community in areas ...
Oct. 9, 2015 03:00 AM EDT Reads: 728
Internet of Things (IoT) will be a hybrid ecosystem of diverse devices and sensors collaborating with operational and enterprise systems to create the next big application. In their session at @ThingsExpo, Bramh Gupta, founder and CEO of robomq.io, and Fred Yatzeck, principal architect leading product development at robomq.io, discussed how choosing the right middleware and integration strategy from the get-go will enable IoT solution developers to adapt and grow with the industry, while at the same time reduce Time to Market (TTM) by using plug and play capabilities offered by a robust IoT ...
Oct. 9, 2015 02:00 AM EDT Reads: 2,220
Today's connected world is moving from devices towards things, what this means is that by using increasingly low cost sensors embedded in devices we can create many new use cases. These span across use cases in cities, vehicles, home, offices, factories, retail environments, worksites, health, logistics, and health. These use cases rely on ubiquitous connectivity and generate massive amounts of data at scale. These technologies enable new business opportunities, ways to optimize and automate, along with new ways to engage with users.
Oct. 9, 2015 02:00 AM EDT Reads: 167
The Internet of Things (IoT) is growing rapidly by extending current technologies, products and networks. By 2020, Cisco estimates there will be 50 billion connected devices. Gartner has forecast revenues of over $300 billion, just to IoT suppliers. Now is the time to figure out how you'll make money – not just create innovative products. With hundreds of new products and companies jumping into the IoT fray every month, there's no shortage of innovation. Despite this, McKinsey/VisionMobile data shows "less than 10 percent of IoT developers are making enough to support a reasonably sized team....
Oct. 9, 2015 02:00 AM EDT Reads: 209
"In the past year we've seen a lot of stabilization of WebRTC. You can now use it in production with a far greater degree of certainty. A lot of the real developments in the past year have been in things like the data channel, which will enable a whole new type of application," explained Peter Dunkley, Technical Director at Acision, in this SYS-CON.tv interview at @ThingsExpo, held Nov 4–6, 2014, at the Santa Clara Convention Center in Santa Clara, CA.
Oct. 9, 2015 01:45 AM EDT Reads: 7,028
Through WebRTC, audio and video communications are being embedded more easily than ever into applications, helping carriers, enterprises and independent software vendors deliver greater functionality to their end users. With today's business world increasingly focused on outcomes, users' growing calls for ease of use, and businesses craving smarter, tighter integration, what's the next step in delivering a richer, more immersive experience? That richer, more fully integrated experience comes about through a Communications Platform as a Service which allows for messaging, screen sharing, video...
Oct. 9, 2015 12:00 AM EDT Reads: 1,135
SYS-CON Events announced today that Dyn, the worldwide leader in Internet Performance, will exhibit at SYS-CON's 17th International Cloud Expo®, which will take place on November 3-5, 2015, at the Santa Clara Convention Center in Santa Clara, CA. Dyn is a cloud-based Internet Performance company. Dyn helps companies monitor, control, and optimize online infrastructure for an exceptional end-user experience. Through a world-class network and unrivaled, objective intelligence into Internet conditions, Dyn ensures traffic gets delivered faster, safer, and more reliably than ever.
Oct. 8, 2015 10:00 PM EDT Reads: 597
The IoT market is on track to hit $7.1 trillion in 2020. The reality is that only a handful of companies are ready for this massive demand. There are a lot of barriers, paint points, traps, and hidden roadblocks. How can we deal with these issues and challenges? The paradigm has changed. Old-style ad-hoc trial-and-error ways will certainly lead you to the dead end. What is mandatory is an overarching and adaptive approach to effectively handle the rapid changes and exponential growth.
Oct. 8, 2015 09:00 PM EDT Reads: 119
Mobile messaging has been a popular communication channel for more than 20 years. Finnish engineer Matti Makkonen invented the idea for SMS (Short Message Service) in 1984, making his vision a reality on December 3, 1992 by sending the first message ("Happy Christmas") from a PC to a cell phone. Since then, the technology has evolved immensely, from both a technology standpoint, and in our everyday uses for it. Originally used for person-to-person (P2P) communication, i.e., Sally sends a text message to Betty – mobile messaging now offers tremendous value to businesses for customer and empl...
Oct. 8, 2015 05:30 PM EDT Reads: 233
Can call centers hang up the phones for good? Intuitive Solutions did. WebRTC enabled this contact center provider to eliminate antiquated telephony and desktop phone infrastructure with a pure web-based solution, allowing them to expand beyond brick-and-mortar confines to a home-based agent model. It also ensured scalability and better service for customers, including MUY! Companies, one of the country's largest franchise restaurant companies with 232 Pizza Hut locations. This is one example of WebRTC adoption today, but the potential is limitless when powered by IoT.
Oct. 8, 2015 04:30 PM EDT Reads: 7,474
You have your devices and your data, but what about the rest of your Internet of Things story? Two popular classes of technologies that nicely handle the Big Data analytics for Internet of Things are Apache Hadoop and NoSQL. Hadoop is designed for parallelizing analytical work across many servers and is ideal for the massive data volumes you create with IoT devices. NoSQL databases such as Apache HBase are ideal for storing and retrieving IoT data as "time series data."
Oct. 8, 2015 02:45 PM EDT Reads: 499


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Vt. state police search for 500-lb steam cookerApril 27,2013MARLBORO Anyone see a missing 500-pound steam cooker?
Vermont State Police are investigating the theft of the cooker, which had been located behind the Skyline restaurant on Route 9 in Marlboro.
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- Report: #542776
Report - Rebuttal - Arbitrate
Complaint Review: Cheap Cars and Trucks, Jerimiah Boss
Cheap Cars and Trucks, Jerimiah Boss6640 N, Government Way Suite A. Coeur d'Alene, Idaho United States of America
Cheap Cars and Trucks, Jerimiah Boss Lemon Car Dilemma, December 2009 I wish to press charges for the following and or a law suit with full refu Scammed me on a van, wont agree to cover the repair cost. Coeur d'Alene Idaho
*General Comment: Co victim responds
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Lemon Car Dilemma, December 2009 I wish to press charges for the following and or a law suit with full refund, or cover the cost of all repairs, including the dealerships at fault for the drivers side dented door.
Thursday December 17th 2009, in the late afternoon bout a used 1995 Plymouth voyager, full tank of gas included and what I thought was all fluids topped off. I get the car home in Lewiston Idaho from CD'A Idaho. And the next day Friday it's leaking trani fluids, as well as on Saturday the engine failed to run after going up a hill and stalling out in the slow lane. Under the lemon law since its not been a full 3 days from the purchase. (Even after signing on a non warranty wavier), does the dealer consume full responsibilities for the repair cost and towing fee to my mechanic of Lewiston. Name of the bushiness as well. I feel as though I have been cheated and shafted. The name of the Dealership with contact info in Coeur d'Alene, ID 83814 is along with the sales person. {Cheap Cars and Trucks Jeremiah Boss general manager; 6640 N. Government Way Suite A. Coeur d'Alene, ID 83814} Cell 208-704-0980, Business phone: 208-772-9977, Fax: 208-772-9970. I wish to have this investigated, and or sue the owner and make him either replace the vehicle; second I get a full return of the vehicle to the dealership and a full refund and or money back the $1500.00 roughly. Three law suit and the repair cost covered in full by the dealership. I want this investigated pronto, before Christmas thanks. This report is also being sent to the better business bureau. I can be reached at the above contact information.
This report was posted on Ripoff Report on 12/20/2009 05:43 PM and is a permanent record located here: http://www.ripoffreport.com/r/Cheap-Cars-and-Trucks-Jerimiah-Boss/Coeur-dAlene-Idaho-83501/Cheap-Cars-and-Trucks-Jerimiah-Boss-Lemon-Car-Dilemma-December-2009-I-wish-to-press-char-542776. The posting time indicated is Arizona local time. Arizona does not observe daylight savings so the post time may be Mountain or Pacific depending on the time of year.
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AUTHOR: Dave - (U.S.A.)
SUBMITTED: Wednesday, February 10, 2010POSTED: Wednesday, February 10, 2010
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Knights Inn East Knoxville
No prices found through our partners. Please contact the business directly or check some of our recommended alternatives.
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- Continental Breakfast
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TripAdvisor Reviews Knights Inn East Knoxville
Travel Blogs from Knoxville
Day 6
Blue ridge parkway in the smoky mountains. Rode a stretch that was so narrow and as tight as anything on the dragon. Kent started the day with his shirt on backwards! Ha!! Cool up in the mountains , had to pull out a jacket! Great day! Heading out tomorrow back to Kentucky starting ...
Longer day than expected
... male labs. He was in LOVE with Bruce... but could of cared less about Missy (a smaller mix female)
5) Dolly Parton was everywhere. She has a highway named after her, a theme park... i think a splash park... it's just dolly dolly dolly.
6) Traffic was awful. Awful.
7) Moonshine from TN is no joke... I'm pretty sure you catch a buzz just from walking past the jar....
The night in TN was great, Phil and his family ...
Lake City FL. To Knoxville TN
Started early. Went to Starbucks then topped off at Supercharger. Stopped to charge at Tifton and Macon GA (where an ICE was in charging space) before arriving in Atlanta about 12:30. Chargers are located in Atlantic Station parking lot. About 20 chargers and half full. Never seen so many Teslas!! Had lunch there then scrambled to get out of Atlanta before traffic. Beautiful day and great time and drive until I pulled ...
Day 24
At our first night at an airbnb, we were made to feel very welcome by Vicki and Buddy, they showed us to a very lovely large bedroom with an onsuite. Having slept very well the next morning we left early for Gatlinburg, the second attempt! No short cuts, no scenic routes just the Interstate all the way. We had to pass through Pigeon Forge and we were amazed by the buildings ( if you can call them that ). A play ground ...
Day 23
... we decided to head for Knoxville.We had booked through Airbnb for tonight and needed to be there at 4pm as Vicki the owner was going out. It was at this point it all went wrong. We travelled through heavy rain,clouds and very twisting roads up and down mountains. We realised afterwards this was Smokey Mountain Pass, reaching a height of over 5000 ft.(No wonder we went through clouds) and realised were going the wrong way when we saw the Cherokee road sign ( this ...


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Page 1
Mapping the structure and conformational movements of
proteins with transition metal ion FRET
Justin W. Taraska1, Michael C. Puljung1, Nelson B. Olivier2, Galen E. Flynn1, and William N.
1 Department of Physiology and Biophysics, Howard Hughes Medical Institute, University of,
Washington, Seattle, Washington 98195
2 Department of Chemistry, Massachusetts Institute of, Technology, Cambridge, Massachusetts
Visualizing conformational dynamics in proteins has been difficult, and the atomic-scale motions
responsible for the behavior of most allosteric proteins are unknown. Here, we report that FRET
between a small fluorescent dye and a nickel ion bound to a di-histidine motif can be used to
monitor small structural rearrangements in proteins. This method provides several key advantages
over classical FRET including the ability to measure the dynamics of close range interactions, the
use of small probes with short linkers, a low orientation dependence, and the ability to add and
remove unique tunable acceptors. We used this 'transition metal ion FRET' approach along with
x-ray crystallography to determine the structural changes of the gating-ring of the mouse
hyperpolarization-activated cyclic nucleotide-regulated ion channel HCN2. Binding of cAMP to
the isolated carboxyl-terminal region of HCN2 caused a structural rearrangement involving a
movement of the C-helix towards the β-roll of the cAMP-binding domain and a movement of the
F′ helix of the C-linker, along with a stabilization of the secondary structure of the helices. Our
results suggest a general model for the conformational switch in the cyclic nucleotide-binding site
of cyclic nucleotide-regulated ion channels.
In allosteric proteins, the binding of a ligand stabilizes conformational rearrangements,
resulting in a switch between inactive and active states. These movements can propagate
long distances to affect activity tens of angstroms from the ligand-binding site. In HCN
channels, the direct binding of cAMP to a cytoplasmic ligand-binding domain stabilizes the
opening conformational changes in the channel pore located more than 50 Å away1.
However, like in most allosteric proteins, the direction, magnitude, and nature of
conformational changes in the channel are unknown.
Fluorescence resonance energy transfer (FRET), in which light energy absorbed by a donor
is transferred to a nearby acceptor, is a powerful tool for measuring changes in molecular
distances2. The efficiency of FRET falls off with the sixth power of the distance between
the two molecules, making FRET exquisitely sensitive to changes in distance. However, as a
Address correspondence to: William N. Zagotta, Ph.D. Dept. of Physiology and Biophysics, Box 357290, University of Washington,
Seattle, WA 98195-7290, Tel: (206) 685-3878, Fax: (206) 543-0934, firstname.lastname@example.com.
J.W.T designed the experiments and analysis. J.W.T and M.C.P. performed fluorescence experiments. N.B.O, G.E.F, and W.N.Z,
performed the crystallography. J.W.T and W.N.Z. wrote the manuscript.
NIH Public Access
Author Manuscript
Nat Methods. Author manuscript; available in PMC 2010 January 1.
Published in final edited form as:
Nat Methods. 2009 July ; 6(7): 532–537. doi:10.1038/nmeth.1341.
NIH-PA Author Manuscript
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consequence of this steep distance dependence, FRET can measure distances effectively
only in a narrow window around R0, the distance at which FRET efficiency is 50%2.
Classical FRET methods are not always well suited to study intramolecular movements in
proteins. This is chiefly due to the long R0 values of most fluorescent dye or protein FRET
pairs (30–60 Å), their large sizes (15–30 Å), and their long flexible attachment linkers (10–
15 Å) 3,4. Combined with the dependence of the FRET on the relative orientation of the
fluorophores, these properties can complicate the interpretation of FRET results. While
lanthanide resonance energy transfer (LRET) methods have less orientation dependence than
FRET, LRET still utilizes large chelating groups and dyes with long linkers, has similarly
long R0 values (~30–100 Å), and has the added complication of very long excited state
What is needed to map conformational rearrangements in proteins is a technique that can
work over shorter distances than classical FRET. We have developed such a method called
"transition metal ion FRET", which has shorter R0 values (about 10 Å), uses small dyes with
short linkers, and is not sensitive to the orientation problems usually associated with FRET.
Unlike NMR, EPR, and crystallography, transition metal ion FRET could theoretically be
applied to proteins of any size, in membranes, on extracellular domains of proteins in living
cells, and potentially on single molecules. It can measure the kinetics of the conformational
change in real time and can be paired with other functional measurements. Here, we utilize
this method along with x-ray crystallography to explore the conformational motions of the
carboxyl terminal region of HCN2 during ligand-activation. Our results demonstrate the
ability of transition metal ion FRET to reveal structural rearrangements in proteins and
provide a model for allosteric movements of HCN2.
Transition metal ion FRET
Our fluorescence method relies on two previous observations. First, transition metal ions
such as Ni2+ and Cu2+ are colored and their weak absorbances overlap with the emission of
fluorescent dyes including fluorescein (Fig. 1a). Thus, Ni2+ and Cu2+ can be used as short-
range FRET acceptors6,7. Indeed, the R0 for Ni2+ and fluorescein (12 Å) is 4 times shorter
than standard FRET pairs such as CFP, YFP (50 Å) or fluorescein, rhodamine (54 Å) 8,9.
The R0 value between Cu2+ and fluorescein is longer (16 Å), allowing transition metal ion
FRET to be tuned to different distance ranges. Second, a metal-binding site can be
engineered into proteins with minimal perturbation to the backbone by introducing two
histidines spaced one turn away on an α-helix (Fig. 1b) 10,11. We reasoned that by
engineering a metal ion-binding motif into α-helices, and by introducing a fluorophore such
as fluorescein onto a cysteine residue, we could use FRET to observe small conformational
rearrangements in proteins. The efficiency of FRET (E), and hence the distance between the
donor and acceptor, can easily be determined by measuring fluorescence before (F) and after
(Fmetal) the addition of metals (E = 1-Fmetal/F). To test the use of transition metal ions as
short-range FRET acceptors, we measured FRET in a set of model α-helical peptides based
on the sequence AAAAKAAAKAAAAAKA. The peptides contained a single reactive
cysteine at the second amino acid position (C2) and a pair of histidines spaced at increasing
distances along one face of the helix (Fig. 1b). We modified the cysteine with the reactive
fluorophore fluorescein-5-maleimide (F5M). We performed fluorescence quenching
experiments on the peptides in solution. In a peptide with a metal binding site adjacent to the
fluorophore (C2H3H6), we observed substantial quenching of the emission spectrum of
fluorescein with increasing concentrations of Ni2+ (Fig. 1c). While 1 mM Ni2+ produced no
appreciable FRET (1-Fmetal/F) in the C2F5M control peptide containing no histidines (3% ±
4 %), FRET efficiency decreased with distance in peptides containing the di-histidine motif:
C2H3H6 (43% ± 4 %); C2H6H10 (27% ± 4%); and C2H10H14 (13% ± 2%) (Fig. 1d). The
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small decrease in fluorescence seen in the C2F5M control at 10 mM Ni2+ was likely due to
collisional quenching of fluorescein by Ni2+ in solution and had a Stern-Volmer quenching
constant (Ksv) of 18 M−1 (Supplementary Fig. 1). These data indicate that Ni2+ can be used
as a close-range distance-dependent acceptor for fluorescein. Cu2+, which has a longer R0
than Ni2+, exhibited more robust distance-dependent FRET with fluorescein-modifed
peptides: C2H3H6 (84% ± 5); C2H6H10 (77% ± 2); and C2H10H14 (50% ± 7%) (Fig. 1e).
To demonstrate that our data could be explained by a FRET mechanism, we compared
distances calculated from FRET to distances predicted from model peptides (Fig. 1f).
Assuming an α-helical structure of the peptide and a modeled position of the fluorophore
and metal binding site, the FRET efficiencies obtained for both Ni2+ and Cu2+ at all three
metal ion binding-sites are fit well by the Forster equation with R0 values of 12 Å and 16 Å
for Ni2+ and Cu2+ respectively. These results support the idea that quenching occurred by a
FRET mechanism.
FRET measurements of the HCN2 holo state
Cyclic nucleotide-binding domains (CNBD) are found in a variety of proteins. The domain
consists of a β-roll and a carboxyl-terminal α-helical region (B- and C-helix), which caps
the cAMP-binding site (Fig. 2a). In each of the subunits of the cyclic nucleotide-regulated
ion channel HCN2, a CNBD is linked through a series of 6 helices (C-linker) to a
transmembrane pore-forming region12. HCN2 channel subunits assemble as a symmetric
tetramer to form an ion-conducting pore and a cytosolic gating ring made from the C-linker
and CNBD from each subunit (Fig. 2b). In cyclic nucleotide-regulated channels, the binding
of cAMP to the CNBD stabilizes conformational rearrangements that propagate up through
the C-linker and allow the pore to open12.
Previous models of how cAMP activates HCN2 channels have suggested that structural
changes in the C-helix and C-linker occur12. To elucidate the structural transitions that
occur in these regions, we mutated, expressed, and purified a cysteine-less version of the
carboxyl-terminal region of HCN2. While a tetramer in the channel, this fragment is a
monomer in solution1. We solved the X-ray crystal structure of this fragment bound to
cAMP (holo state). The structure was virtually identical to that of the wild-type holo state
HCN2 (RMSD of main chain atoms is 0.7 Å) (Fig. 2a,b and Supplementary Table 1)1. We
then introduced single cysteines at surface-exposed sites in the F′-helix of the C-linker
(N520C) or in the β-roll of the CNBD (K570C) and covalently modified them with
fluorescein-5-maleimide. We introduced Ni2+-binding motifs (His-X3-His) at 4 positions
along the C-helix: M620H, A624H; A624H, V628H; T627H, D631H; and I630H, D634H.
To map distances in the holo state, we measured the fluorescence with increasing
concentrations of Ni2+ (Fig. 2c). All measurements were done at saturating concentration of
cAMP (250 μM) (Supplementary Fig. 2). Constructs labeled at N520C in the C-linker
showed robust concentration-dependent FRET that plateaued at about 500 μM Ni2+ (Fig.
2c). Notably, at 500 μM Ni2+, each metal-binding site exhibited a different amount of
(74.9% ± 1.2%); and I630H, D634H (43.5% ± 2.3%) (Fig. 2c). Constructs labeled at K570C
in the β-roll also produced significant FRET values at saturating Ni2+: M620H, A624H
(30.3% ± 1.9%); A624H, V628H (29.4% ± 1.7%); T627H, D631H (37.4% ± 1.6%); and
I630H, D634H (36.2% ± 1.8%) (Fig. 2d).
Unexpectedly, the control channel construct N520CF5M which contains no engineered Ni2+-
binding motifs also showed Ni2+-dependent FRET (27.2% ± 3.5%) (Fig. 2c). Cu2+ produced
a larger amount of FRET in the control N520CF5M (Supplementary Fig. 3)6,7. The large
Cu2+-dependent FRET signal in the control barred the use of Cu2+ to map distances in
HCN2. However, these data are consistent with the background quenching being the result
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of FRET between a native metal-ion binding site in HCN2 and N520CF5M. While the Ni2+
effect was smaller and somewhat lower affinity (44 μM) than that of the constructs with
engineered Ni2+-binding motifs, it was still higher affinity than that of the control peptide
containing no histidines (C2F5M ) (>50 mM) (Fig. 1c and Supplementary Fig. 1). These
results suggest that the HCN2 carboxyl-terminal region contains a native metal ion-binding
site near the F′-helix. In the K570CF5M control channel fragment, the native Ni2+-binding
site produced less FRET (18.2% ± 2.6%), indicating that the native Ni2+ site is farther from
K570CF5M than N520CF5M (Fig. 2d).
To relate the holo state structure of the HCN2 carboxyl-terminal fragment in solution to the
crystal structure, we compared distances determined from FRET to distances determined
from X-ray crystallography. We calculated distances from FRET efficiencies using a Forster
equation that took into account FRET from the native Ni2+ binding sites and assumed
random orientation of the donor and acceptor. This is a reasonable assumption given the
anisotropy of the fluorophores (N520CF5M 0.216±0.001, K570CF5M 0.178±0.002)
(Supplementary Fig. 4) and the multiple absorption dipoles of the Ni2+ ions13,14. To
separate the specific FRET signal of the engineered di-histidine sites from the native Ni2+
binding site we used a version of the Forster equation that considers FRET to multiple
acceptors and held the distance to the native site constant15–18. While we do not know if
there is one or multiple native metal sites, a single apparent distance term could be used to
correct FRET signals15–18. Using spatial coordinates from the holo HCN2 crystal structure,
we modeled the Ni2+-binding motifs into constrained positions along the C-helix.
Fluorophore positions (modeled as a point source) that fit the experimental FRET distances
were found using a best-fit least-squares analysis. Figure 2e and Supplementary Figure 5
shows one possible position of the dye that overlapped the point source. The modeled
distances closely match our experimental values (Fig. 2f) with a RMSD of 2 Å. The single
outlier likely resulted from difficulties in measuring distances far from R0 Movements of
excited state fluorophores or a distribution of fluorophore positions can bias FRET
measurements towards shorter distances or R0 respectively9. From this analysis combined
with our measurements on peptides (Fig. 1) we conclude that the structure in solution
closely matches the structure in the crystal, and that our FRET method can map spatial
positions within proteins.
X-ray crystal structure of the HCN2 apo state
To understand how cAMP-binding changes the conformation of HCN2, we solved the x-ray
crystal structure of the wild-type HCN2 carboxyl-terminal fragment in the absence of cAMP
(apo state) (Fig. 3a,b and Supplementary Table 1). Surprisingly, the overall structure is very
similar to that of the holo state (Fig. 3c). The first 5 helices of the C-linker as well as the β-
roll are virtually identical (RMSD 0.5 Å). We could resolve several bromide ions in the
structure with two located in the cyclic nucleotide binding pocket, one of which occupies the
position equivalent to the phosphate group of cAMP in the holo state structure
(Supplementary Fig. 6). We observed two major differences between the holo and apo state
structures. First, the F′-helix adopts a less helical structure in the apo state (Fig. 3c,d).
Second, the carboxyl-terminal portion of the C-helix was no longer resolved, indicating that
the end of the C-helix becomes less ordered. This lack of structural information of the
carboxyl-terminal portion of the C-helix precludes us from determining the precise
conformation of the C-helix in the apo state via crystallography. Furthermore, the small α-
helical amino-terminal portion of the C-helix visible in the apo-state crystal, which was
nearly identical to the holo state structure, was involved in crystal contacts between
tetramers and therefore might not accurately represent the solution structure of this region
(Supplementary Fig. 7). Nevertheless, these data suggest that cAMP-dependent
rearrangements occur in the F′- and C-helices.
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FRET measurements of cAMP-induced structural changes
Because we were unable to fully determine the rearrangements of the C-helix from
crystallography, we turned to our transition metal ion FRET method. The Ni2+-dependence
of FRET was measured for each of the previous constructs, this time without cAMP (Fig.
4a,b). The control construct containing N520CF5M in the F′-helix and no engineered Ni2+-
binding motifs exhibited a small decrease in FRET in the absence of cAMP, consistent with
a small change in distance between this position and the native Ni2+-binding site (Fig. 4c).
We did not observe this change with K570CF5M, which indicates that, while the native Ni2+
site is stationary, the F′-helix (N520CF5M) makes a small movement during cAMP binding.
These data are consistent with both the crystallography data and previous studies of CNGA1
channels19. We observed changes in FRET for all of the N520CF5M and K570CF5M
constructs containing engineered Ni2+-binding sites in the C-helix (Fig. 4c). To translate
FRET changes into distances, we calculated distance changes (holo vs. apo) according to the
Forster equation taking into account FRET from the native Ni2+-binding site15–18. We
detected large movements between both N520CF5M and K570CF5M and the first and second
pair of histidines in the C-helix (Fig. 4c). In contrast, the middle and end of the C-helix
(T627H, D631H and I630H, D634H) showed only small changes in distance. Since these
movements were also detected with the fluorophore at K570C, which undergoes little or no
movement itself, it seems highly likely that the FRET changes we observed arise primarily
from movements of the C-helix and not movements of the fluorophores themselves. At
every position, FRET increased upon cAMP-binding, which suggests that the C-helix,
particularly the amino-terminal end of the C-helix, translates toward the β-roll of the CNBD.
Along with distance measurements, our FRET method provides information about protein
secondary structure. The affinity of di-histidine motifs for transition metal ions is sensitive
to helix stability10,11,20. Indeed, while individual metal binding sites within the same
proteins can have diverse Kd's, helix stability tends to increases the affinity of engineered
sites for metal11,20,21. In HCN2, cAMP-binding caused a significant increase in the
affinity for Ni2+ at each introduced Ni2+-binding site (but not the native sites) in N520CF5M
(P < 0.05, student's t-test) (Fig. 5a and Supplementary Table 2). We observed a similar
increase in affinity for proteins labeled at K570CF5M (Fig. 5b and Supplementary Table 2).
The large shifts in Ni2+ affinity correspond to Δ Δ G values between − 0.4 and − 1.7 kcal/
mol, values similar to the energetics of helix stabilization observed in other
systems11,20,21. The observed shifts indicate that cAMP-binding stabilizes the α-helical
structure of the C-helix. These data demonstrate the ability of metal ion FRET to reveal
secondary structure transitions in proteins.
Until now, models of CNBD dynamics suggested that the C-helix makes a rigid-body
movement to cap the CNBD12,22. Crystal structures of CNBD-containing proteins
including protein kinase A (PKA), exchange protein activated by cAMP (EPAC), and the
bacterial cyclic nucleotide regulated channel M. loti K have shown movements of the C-
helix in response to cAMP-binding23–25. Each of these structures shows different and
sometimes large conformational rearrangements. In the M. Loti K channel alone, x-ray
crystallography revealed six distinct conformations of the C-helix, making it difficult to
infer how the C-helix actually moves in solution23. Clearly, it is sometimes difficult to
determine the conformational motions of a protein from crystal structures alone.
For HCN2, crystallography did not reveal substantial differences in the resolved portion of
the C-helix between the holo and apo states. We believe that crystal contacts in this region
likely stabilized the C-helix in a non-physiological conformation. With transition metal ion
FRET, however, we measured cAMP-induced rearrangements of the C-helix in solution, an
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text:
Pocono Farms is just north of Mt Pocono and is a superb parkland style course designed by Art Wall.
The layout is scenic with tight tree lined holes over rolling and sloping fairways with little room off the tee. There's a good amount of sand in play, although mostly greenside, a few holes along Echo Lake, otherwise a couple of holes with water in play. It's rather short at 6,100 yards, 125 slope from the tips with two shorter tee boxes.
No golf course reviews have been submitted on this course.


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Date: 1900 ca.
Description: Cary Bass, standing next to his Great Uncle Cary, who is seated. They are most likely seated inside the Bass family home, 207 Clay Street.
Description: Crowd gathered at the Montello railroad station.
Date: 1925
Description: View down dirt road along the shoreline of Crystal Lake. A man and a child stand under a tree's shade, near a pier where a man is seated on the dock. There...
Date: 1910 ca.
Description: A workman poses beside a belt-driven polisher inside a shed at the Montello granite quarry. Two other men relax in the background near an open door where n...
Description: Great Uncle Cary posing with Everetta and Cary Bass. They are most likely seated inside the Bass family home, 207 Clay Street.
Description: Girls from a Sunday school class. Everetta, the photographer's daughter, stands on right with hand warmer (muff).
Description: View from shoreline of two women standing on a dock on Crystal Lake. Just beyond the dock, a man and woman fish from a boat.
Description: Two men stand on a ledge in the granite quarry in Montello. Three other men stand below. The side of the quarry rises up above the group in the background....
Description: Family group resting on Court House hill, near Perkin's cottage.
Date: 1908-07
Description: Workmen with teams of horses excavate for the new Mercantile Store, later the M.M. Smart Store. A woman is walking on the street nearby, and a boy or man w...
Description: Sunday school class giving a patriotic presentation.
Date: 1905 ca.
Description: A view of the Cramer residence, a summer home of Mrs. William Cramer.
Description: Exterior view of front of the Montello House hotel.
Date: 1915 ca.
Description: A church in Westfield.
Description: Ada Bass with her son, Edward Cary, sitting on a couch.
Description: View of the Montello granite quarry grounds shows piles of granite paving blocks. In the background, a team of horses pulls a wagon under an elevated narro...
Date: 1927
Description: Two women pose outside their lakefront cabin.
Description: Large group of people at a picnic at Kratz Grove on the south shore of Buffalo Lake.
Date: 1926-06
Description: A man poses holding a fish in both hands. Two boys wait at the service window of a log cabin.
Date: 1934-02-27
Description: CWA road crew leveling a grade on a county road near Westfield. The CWA undertook many road and utility projects during the Great Depression in order to p...
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also noticed when I activate the LOCK on my fob the lights don't blink on my car. ONLY when I UNLOCK it blinks. Does that have anything to do with my alarm going off? I hate to leave it unlocked. It goes off in the middle of the night/wee hrs off the morning, anytime it wants. HELP!!!!
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Enter your login details for Case Studies in Engineering Failure Analysis below. If you do not already have an account you will need to register here.
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Failure analysis of a motor-car coil spring
Available online 28 December 2013
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Failure analysis of a high strength low alloy 0.15C–1.25Cr–1Mo–0.25V steel pressure vessel
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text:
Hwy. 90 / River Road highway in our future?
October 29, 2010 at 9:19 am | Mobile Reader | Print
At the meeting, the Department of Transportation and Development was taking an aggressive approach to a new highway, Tastet said. There is a lot of communication going on.
Currently, Barton Avenue serves as a major connection between Highway 90 and River Road, but it runs through a residential neighborhood. The road has become heavily congested because of the lack of a signal light at the intersection of Barton and River Road, which forces commuters to wait until multi-directional traffic clears before they can turn.
Two railroad tracks also run through the street.
The commission is looking at three options when it comes to building a new highway, Tastet said. One route would call for the extension of Willowdale Boulevard to River Road, while another would be a more easterly route that would run near the Davis Pond Diversion.
The third option, which appears unlikely, would be further west near Barton Avenue.
Everyone was very receptive to the idea of building a new highway, Tastet said. Right now, Barton is so congested and its causing a lot of problems. Ever since I got into office I have received a ton of complaints about it.
Commute time on Barton can last anywhere from 10 to 20 minutes at some times of the day, and it creates a hassle for those that live on the street and have trouble even backing out of their driveways.
Tastet and St. Pierre will visit with the Regional Planning Commission early next month and will go over all three options. Tastet said that the Davis Pond Diversion route would make the most sense.
Rathborne Land Co. owns a majority of the land that such a route would travel through. Tastet said that the company has shown an interest in donating the land needed for the highway.
An extension of Willowdale Boulevard would travel through land owned by Levert Land Co. Jim Hooper, of Levert, told the council in March that the company would consider donating that land to St. Charles Parish, assuming the study shows that a new highway would be feasible.
A new highway would be major because not only would it take pressure off of Barton, but it would also take pressure off of Paul Maillard Road, Tastet said. If a new highway is constructed, Barton would then become a parish street, which is what it should be since it runs through a neighborhood.
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text:
Mark EpsteinIt will hold you, as it held me, in its thrall.
Can Love Last?: The Fate of Romance over Timeby Stephen A. Mitchell
View All Available Formats & Editions
"A beautiful and brilliant reexamination of love and its perils."—Barbara Fisher, Boston GlobeSee more details below
"A beautiful and brilliant reexamination of love and its perils."—Barbara Fisher, Boston Globe
Editorial Reviews
Andrew SolomonThose who read this book will love more wisely because of it.
Barbara FisherA beautiful and brilliant reexamination of love and its perils.
Boston Globe
JoAnn Gutin[T]houghtful,compassionate,and profoundly optimistic.
Judith ShulevitzCheerful,open,and humaneyou'd definitely have wanted him as your analyst.
New York Times Book Review
Publishers WeeklyWhen New York University professor and popular psychoanalysis theorist Mitchell died in December 2000, he left behind a robust body of work that made Freudian theories accessible to all. It's not surprising, then, that this postmortem work should have broad appeal. A combination of clinical case studies, psychoanalytical thought and practical advice, Mitchell's riff on the fragility and necessity of romantic love is written with warmth and intelligence. He manages to simplify some of Freud's most complex theories and give them new significance for those who wonder why love is often a battlefield. Real-life examples, taken from his practice, are an invaluable addition. In a section on guilt, for example, he briefly describes how Freud considered the emotion to be "the linchpin of our ascent from the bestial to the civilized," then brings in the work of Viennese-born analyst Melanie Klein and concludes with the story of "Will," whose tendency toward feeling guilty created havoc with his romantic relationships. By mixing the case study method, so common in self-help books, with scholarly insight, Mitchell creates a work on romance that is rich and multilayered, giving the individual stories more weight and the intellectual commentary more humanity. In his conclusion, Mitchell writes like a loving father penning a wedding day message to his child, gently advising that romance isn't about "a labored struggle to contrive novelty," but instead about tolerance and understanding. It's common advice, but given the rest of the work's depth, humor and rigor, these familiar words take on new, and much welcomed, meaning. (Feb. 14) Forecast: Mitchell was always adept at user-friendly writing, and this work follows in that tradition. Can Love Last? would do well on its own merit, and the Valentine's Day pub date should push sales further. Copyright 2001 Cahners Business Information.
Library JournalThis rather dry academic treatise on romantic love was written by a well-regarded clinician and supervising analyst at the William Alanson White Institute. Mitchell, who died in 2000, was the author of several books including Freud and Beyond, which he wrote with his wife Margaret Black. Popular culture would have us believe that the combination of romance and true lasting love is an oxymoron. Mitchell examines the tension between the ideas of love and romance, and shows how sexuality, illusion, aggression, guilt, control, and commitment interact to contribute to this tension. He also describes the different risks involved in both stable and new love. While drawing heavily on Freud, he examines romance and love from many different psychoanalytic viewpoints. Cases from his practice are described to illustrate his discussion. The scenario of the cocktail party toward the end of the book is especially clear in conveying different theories of consciousness. Recommended for academic and large public libraries. Margaret Cardwell, Christian Brothers Univ. Lib., Memphis Copyright 2002 Cahners Business Information.
Product Details
- 9780393078480
- Publisher:
- Norton, W. W. & Company, Inc.
- Publication date:
- 02/17/2003
- Sold by:
- Barnes & Noble
- Format:
- NOOK Book
- Pages:
- 224
- Sales rank:
- 452,751
- File size:
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