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- AgriPOWER opens doors
- Value from the people
- 2015 County Farm Bureau Presidents Trip to D.C.
- Farm Bureau supports new nutrient bill
- Ohio Farm Bureau's State Priority Issues for 2015
Year in Review: Telling our story
Crawford Countys Greg and Rose Hartschuh became the faces of the new I am Farm Bureau campaign this year aimed at dispelling myths of the organization being big agriculture, and providing a way for Farm Bureau members to stand up, speak out, correct information and tell the stories of who they really are.
I am Farm Bureau could be seen on pins, hats and T-shirts worn by members at local and state Farm Bureau events throughout the year, and it served as the theme for its annual meeting and leadership conference.
The Hartschuhs won an online I am Farm Bureau contest featuring member narratives and photos showcasing the diversity and grassroots nature of Farm Bureau. The grain and dairy farming couple are the fourth generation to work on the farm. Their entry detailed how they make time for each other while managing the farm.
Farm Bureau members range from gardeners and food and wine enthusiasts, to nature lovers and conservationists, fruit and vegetable growers to livestock and grain producers, and everything in between.
View the Hartschuhs entry and those of other finalists of the 'I am Farm Bureau' contest


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My Freemasonry | Masonic Education and Discussion Forum
Brother Tadasu HayashiVariously described as a great, brilliant Diplomat and Statesman, an Anglophile, a world famous figure, patriotic, and a man who lived up to the noble teachings of Freemasonry, Tadasu Hayashi served Japan with uncommon devotion and dedication throughout his life.
He was born in Shomofussa Province (now a part of Chiba Prefecture) on February 22, 1850 into the Sato family and given the boyhood name of Shingoro. His father was a progressive physician who practiced and taught Dutch (western) medicine. At an early age he was adopted into the Hayashi family who lived in Edo (now Tokyo) and were the heredity physicians to the Tokugawa Shogunate. His surname thus became Hayashi and his first name was changed to Tadasu.
At the age of twelve Hayashi was sent to Yokohama to study English under Dr. James C. Hepburn, an American medical missionary, who was credited with romanizing the Japanese language, translating the Bible into Japanese, and compiling the first Japanese-English dictionary. Dr. Hepburn was alluded to by the Japanese as "Kunshi," a term signifying a superior person, and Mrs. Hepburn was responsible for establishing the first co-educational school in Japan. The Hepburns considered Hayashi one of the family and through them he received a thorough foundation in the English language.
In 1866--thirteen years after Commodore Matthew C. Perry (a member of Holland Lodge No. 8 in New York City) had made his first visit to Japan—Hayashi was among the fourteen youths selected by the Tokugawa Shogunate to study the English language, manners and customs in London. At first the students lived in a lodging house, but Hayashi disliked this arrangement (recalling his experience with the Hepburns) since all of them tended to speak only Japanese except when they were attending the University College School. He petitioned to have the group separated and live with private families which was granted.
In 1868 the Tokugawa Shogunate was in process...
Famous Prince Hall Freemasons (2015)In March 6, 1775, Prince Hall and 14 men of color were made masons in Lodge #441 of the Irish Registry attached to the 38th British Foot Infantry at Castle William Island in Boston Harbor, Massachusetts. It marked the first time that Black men were made masons in America.
About a year later, since the conflict between England and America had commenced, the British Foot Infantry left Boston, along with its lodge, leaving Prince Hall and his associates without a lodge. Before the lodge left, Worshipful Master Batt, gave them a "permit" to meet as a lodge and bury their dead in manner and form. This permit, however, did not allow them to do any "masonic work" or to take in any new members.
Under it, African Lodge was organized on July 3, 1776, with Prince Hall as the worshipful master. It wasn't long before this lodge received an additional "permit" from Provincial Grand Master John Rowe to walk in procession on St. John's Day.
On March 2, 1784, African Lodge #1 petitioned the Grand Lodge of England, the Premier or Mother Grand Lodge of the world, for a warrant (or charter), to organize a regular masonic lodge, with all the rights and privileges thereunto prescribed.
The Grand Lodge of England issued a charter on September 29, 1784 to African Lodge #459, the first lodge of Blacks in America.
African Lodge #459 grew and prospered to such a degree that Worshipful Master Prince Hall was appointed a Provincial Grand Master, in 1791, and out of this grew the first Black Provincial Grand Lodge.
In 1797 he organized a lodge in Philadelphia and one in Rhode Island. These lodges were designated to work under the charter of African Lodge #459.
In December 1808, one year after the death of Prince Hall, African Lodge #459 (Boston), African Lodge #459 (Philadelphia) and Hiram Lodge #3 (Providence) met in a general assembly of the craft and organized African Grand Lodge (sometime referred to as African Grand Lodge #I).
In 1847, out of...
Hidden Lessons From Ritual WorkSo many times we seem to look at Ritual Work as not being that important, and that it doesn't have to be done that well. We feel that just need to have more Masons for our Lodges. If we fail to share the teachings properly, who do you think looses?
I would like to have you think back to that first night, it could of been a warm or cool night, that we all share. That night we were so apprehensive , or for the sake of better words, confused as to what was going to happen. Those first words you heard said at the Lodge door, asking questions and wanting answers of you, and how you were treated was only the start of your Masonic life. That life that leads most of us on a continuous journey of Masonic travel the rest of our lives.
For some of us, who had to memorize the Degree and Obligations, we share something that no one else can understand. This task of learning them, that we choose to do, and we did. This struggle teaches us what we all can do with hard work and a true desire to accomplish things.
So many times I have listened to the lectures, and still I find them as interesting as the first time I heard them. Every time I hear them, I find a new perspective that I have missed before. I fear for those that do not choose to listen to them in this way, as they will never find the lessons that are taught there in Masonry.
And as for those that give those grand lectures, they learn as well. They learn how it feels to give that perfect lecture and also when they don't get it perfectly right! Most of the time just stopping for a moment to think or to taking a breath. I think we can all learn hidden lessons here too. How we should overlook everyone's little mistakes that we all make in life. Also to remember sometimes the best intentions go wrong by accident.
"Value Your Word"
For What Worth Hath a Liar!
Source: Bro. Micheal Mayer
The "Model Master"The Model Master
By Most W. Bro William Mercer Wilson
To become the model Master of a Lodge should be the ambition of every Brother, and to discharge with efficiency and zeal the duties of that important office should be his most anxious desire. These duties are not confined to the mere repetition of a few phrases, learned by rote, but he should be enabled to instruct the Craft, not only as to the meaning and origin of our ceremonies, but also to explain to them the philosophy which is veiled in its allegories and illustrated by its symbols.
He should be able, also, to convince his Brethren, that all science and art, legitimately directed, are but lines that radiate towards the great " I AM;" that the Sciences are the media by which we are led to contemplate the goodness, greatness, wisdom and power, of the Great Architect of the Universe; and that the Arts are the modes we have developed of expressing our sense and admiration of the wondrous glories of an Almighty Father which are scattered around us.
The Master of a Lodge should also, in his life and in his conversation, be a model for his Brethren to admire and imitate, and should himself practice, out of the Lodge, those great moral doctrines and virtues which he inculcates within its walls. He should be punctual and methodical in all things, and, both by his character and conduct, command the respect, the esteem, and good will of all men; for, as the Master is supreme in his Lodge, and distinguished by his position in the Craft, so should he also be distinguished as the possessor of an irreproachable character, a dignified demeanor, an expanded intellect, and a liberal education. Happy and prosperous must those Lodges be which are governed by such men! - their time of meeting is looked forward to by the Brethren with the most pleasing anticipation. Prompt at the hour, every Brother is at his station, and the work is carried on with pleasure and profit. The Worshipful...
Mouth to EarBy Brother Ernest Borgnine, 33˚
In 1946, I travelled with a friend down to a little town called Abingdon, Virginia, to see what the Barter Theatre had to offer. It offered nothing except hard work and board. My friend, not accepting the work they offered him, stayed one day - I stayed five years. In that time I grew to love the town and all it offered. The people, in particular, were simply marvellous.
Occasionally I would be assigned to go down to the printing shop and get posters made for the upcoming shows at the Barter Theatre. One day, in talking to the owner of the print shop, one Elmo Vaughan, I found that he belonged to the local Masonic Lodge, No. 48, in Abingdon. My father was also a Mason and had advanced to the Thirty-second Degree in Scottish Rite Masonry, and I told this to Elmo. He was pleased, and sensing his pleasure, I asked him if maybe I could join. He said nothing, continuing his work, and a short while later, I took my posters and left.
The next time I saw Elmo, I asked him again about joining the Masonic Order - again he said nothing - and again my work took me away. We became good friends and finally one day I passed by and again I asked if I could join the Masons. Instantly, he whipped out an application and I hurriedly filled it out. I didn't learn 'til later, that in those days, you had to ask three times.
I was thrilled! Not only was I going to be the first actor ever in Lodge No. 48, but I could just imagine my father's surprise when I would spring the old greetings on him! I wanted only to surprise my Dad - and was I surprised, when after I was made an Entered Apprentice, I found I had to remember everything that happened to me at that event and come back and answer questions about it!
I was assigned to a dear old man of about 92 years of age who, I felt, must have been there when the Lodge first started. He was really of the old school - and he started me out with the foot-to-foot, knee-to-knee and...
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Keller @ Large: Another Exercise In Character Assassination
An ad from Mitt Romney's campaign claims the Obama Administration is out to destroy a signature bipartisan accomplishment of the 90's.
Opinion: 'Retroactively Retired', A Laugher Even By Washington Standards
The demand for W. Mitt Romney to release his tax returns has become bipartisan. Conservative elected officials and pundits know that Romney is taking on so much water over his secrecy and questions of his honesty and that he has no other choice.
Opinion: The Second Coming Of W
Down to the trademark "W", W. Mitt Romney is literally the second coming of the George W. Bush: the economic strategy that led to the worst American economy since the Great Depression and the Bush legacy of secrecy in the face of questions from the American public.
Brown, Warren Wage E-Mail War
Republican U.S. Sen. Scott Brown and his chief Democratic rival, Elizabeth Warren, have vowed to fight attack ads on television, radio and the Internet, but in their emailed appeals to supporters, the two routinely portray each other in the harshest light possible.
Senator Brown Transfers To Maryland National Guard Unit
Senator Scott Brown requested the move and is now working in a major position at the National Guard's Pentagon office.
Mitt Romney Wins Illinois Primary
Front-runner Mitt Romney won the Illinois primary with ease Tuesday night, routing Rick Santorum.
Romney, Santorum Swap Charges In Fiery 20th Debate
Primed for a fight, Mitt Romney and Rick Santorum traded fiery accusations about health care, spending earmarks and federal bailouts.
Romney Admits His Tax Rate Is Lower Than Most Americans'
At a campaign event on Tuesday, Romney confirmed speculation that his tax rate is lower than that of most Americans.
Romney And Other GOP Hopefuls Gear Up For Weekend Debates
Mitt Romney is far ahead in New Hampshire, but the former Massachusetts governor and the other Republican presidential candidates are facing two debates that could help define the contest.
Huntsman Says Time Working Against Him In New Hampshire
Republican presidential candidate Jon Huntsman also announced that the he has won the endorsement of the Boston Globe.


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Jonny's 2 bedroom East Putney London flat is the perfect place to stay. Just a little outside the tourist packed center, it is easily reached by the District Line and is very close to grocery stores, pubs and restaurants... a short walk to the river front, steps away from the Wandsworth Museum.
The flat was spotless, tastefully decorated and very spacious. we made use of the kitchen, his laundry machine, a phone that we could use for local calls on weekends, and limitless incoming calls. Plus we enjoyed catching up with Downton Abbey on his Apple TV.
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When we booked the house for our trip to North Yorkshire, we though that we know what will expect us. When we arrived at the mansion, we were impressed ho much bigger and more luxurious the house is than we expected.
The place is probably the most impressing house I've ever been. Me and my four friends enjoyed the stay at the house very much. During the day we organised some trips to locations like York, Robin Hood's Bay etc. and in the evening we enjoyed the quiet garden of the house.
If you go with a group of friends to have a nice weekend, this is the place to stay.The house offers everything you need and much more.
The host was very flexible with the times to checkin and checkout. We felt very comfortable with him and he made us feel welcome at the house.
Jonny is a very nice host.
He gave us his disponibility and informations we needed to enjoy our holidays in London.
The Apartment is efficient, in forniture, confortabile beds and nice bathroom ad well.
We also used washmachine.
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I picked this house as a place for a get-together with a group of old friends that I met at University and was in no way disappointed! The house, gardens, village of Kirby Overblow, and general setting of North Yorkshire are all absolutely gorgeous; with the house being the real star of the show for our stay.
All of the bedrooms were incredibly comfy and the decor and quality of furnishings/bathrooms/kitchens/lounge were all finished to a very high standard. We enjoyed cooking for each other in the huge kitchen and soaking up the sun on the patios outside.
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Bill Cosby and his attorneys are planning to grill Playboy Mansion's owner Hugh Hefner in order to take down Coby's latest sexual assault accuser.
Playboy boss Hugh Hefner has responded to his longtime friend Bill Cosby's sexual abuse allegations, calling it 'truly saddening'.
Margot Robbie has politely declined Hugh Hefner`s offer to pose naked for Playboy magazine.
Playboy mogul Hugh Hefner`s wife Crystal Harris said they had a mellow wedding anniversary celebration with friends at their house.
David Dobkin is in talks to take charge of a movie biopic based on the life of Playboy founder Hugh Hefner.
Model Crystal Harris says she and her husband Hugh Hefner have a strong marriage.
Playboy mogul Hugh Hefner`s marriage with Crystal Harris is reportedly on the rocks.
Helen Flanagan will reportedly meet Playboy owner Hugh Hefner next month to discuss the possibility of her posing for the magazine.
Hugh Hefner and his wife Crystal Harris got into the Halloween spirit by dressing up as Miley Cyrus and Robin Thicke from their controversial MTV VMA performance.
Farrah Abraham may not be what Hugh Hefner and his team are looking for, as they have already rejected her pleas to appear in the magazine twice.
Hugh Hefner`s former girlfriend, Holly Madison has got engaged to her boyfriend Pasquale Rotella, who is also the father of her three-month-old daughter Rainbow Aurora.
Hugh Hefner and his wife Crystal Harris have purchased an architectural home in the Hollywood Hills for USD 4.9 million.
Crystal Hefner`s parents are not only supportive of her relationship with Playboy mogul Hugh Hefner , but they also love her lavish lifestyle.
Hugh Hefner, who turned 87 on Tuesday, isn`t showing signs of slowing down.
Hugh Hefner has admitted that he has slept with more than a thousand women, but insisted that he never cheated in any of his previous two marriages.
Hugh Hefner`s wife Crystal has proved that she has more than just good looks, as she showed off her deejay skills at Playboy`s annual Mansion Masquerade party on Saturday.
Police rushed to the Playboy mansion recently in response to a bogus 911 call reporting an assault inside the home.
Playboy boss Hugh Hefner`s young wife Crystal Harris has put her wedding dress up for auction.
Playboy mogul Hugh Hefner has paid tribute to his longtime assistant following her death.
Playboy mogul Hugh Hefner and Crystal Harris signed an "ironclad" prenup before they tied the knot on December 31, according to sources.
Day in Pics: 30th March
Showbiz World - March 30, 2015
Katrina Kaif's wax statue at Madame Tussauds unveiled!


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Loyola's online tutorial, a requirement for next year's incoming class, is delivered as an animated cartoon that introduces students to a core value of the college-the importance of aca…
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Whether they're downloading music, videos, and software or updating their profile on social networking sites, students are bombarded by a host of online ethical issues. Clarkson has created an…


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science metal prints, research metal prints, medical metal prints, medicine metal prints, forensic metal prints, field metal prints, study metal prints, legal system metal prints, court of law metal prints, crime metal prints
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Bring your artwork to life with the stylish lines and added depth of a metal print. Your image gets printed directly onto a sheet of 1/16" thick aluminum.
The aluminum sheet is offset from the wall by a 3/4" thick wooden frame which is attached to the back. The wooden frame includes a hanging wire for easy mounting on your wall (see photo on the left).
All metal prints ship within 3 - 4 business days and arrive "ready to hang" with mounting hooks and nails.
Metal prints are extremely durable. They're lightweight. They won't bend, and they're water resistant.
The high gloss of the aluminum sheet complements the rich colors of any image to produce stunning results.
For additional product photos and information, visit our metal prints product page.
The precise apparatus of metric photography. A photograph from Alphonse Bertillon's photo album from his exhibition at the 1893 World's Columbian Exposition in Chicago. Alphonse Bertillon (1853-1914) was a French police officer and biometrics researcher who created anthropometry, an identification system based on physical measurements. Anthropometry was the first scientific system used by police to identify criminals. In this system the person was identified by measurement of the head and body, individual markings (tattoos, scars) and personality characteristics. The measurements were made into a formula that would apply to only one person and would not change. The system was eventually found to be flawed, however, because often two...
Science Source� Images offers rights managed and royalty free licensing for a comprehensive library of stock imagery. Our content is carefully selected and accurately captioned to satisfy the needs of creative professions like you! Science Source� is a registered trademark of Photo Researchers, Inc.
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Always Thankful? - Today's Christian Videos
I'll Have a Bluegrass Christmas Without Shoes - @church-ny-fulton-first-united-methodist-church
00:02:59 853 views
Embed church-ny-fulton-first-united-methodist-church 318 views 2010-10-12T16:21:03 Reverend Mark Kimpland of the Fulton First United Methodist Church (www.fultonfumc.org) uses the example of Jesus and the lepers to teach the young people about being thankful for what we have. You can see more on our website. Please visit us today and say helo.
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Rothamsted Insect Survey
50 Years of Rothamsted Insect Survey
Rothamsted is proud to celebrate 50 years of the Insect Survey. The data are valuable for a wide range of fundamental, strategic and applied studies. The aphid bulletin gives weekly counts of aphids throughout the UK.
Department Link
This page is part of the Agroecology department
The AgroEcology group aims to understand the ecological mechanisms that deliver sustainable crop production. We have expertise in movement and spatial ecology of pests and pollinators, above and below-ground functional biodiversity and weed ecology.
Vacancies, employment
The Rothamsted Insect Survey (RIS) is a BBSRC-supported National Capability, attracting additional funding from Research Councils, crop levy boards, the agrochemical industry and others. We are seeking an entomologist with expertise in molecular techniques to play a leading role in the exploitation of the RIS long-term collection of specimens from the UK suction-trap network, and current specimens from the suction-trap and light-trap networks.
Details of all external vacancies will be posted here as they become available. If you find nothing that interests you this visit, please call again. Unfortunately we do not have the resources to hold CVs against possible future vacancies. We can only accept applications quoting a relevant and appropriate vacancy number.
An Equal Opportunities Employer
As users of the disability symbol, Rothamsted Research guarantees to interview all disabled applicants who meet the minimum criteria for our vacancies. For more information click here.
We will aim to provide related internet links but Rothamsted cannot be responsible for their content.


text: Images Screenshot of the online home page of Sony Corporation of America. AIBO entertainment robot, model ERS-111. Former employees of a Sony plant in the French village of Pontonx-sur-l'Adour protest in front of the factory in April 2009 by symbolically planting crosses that bear the names of workers whose jobs had been eliminated. Smoke continues to rise from a Sony Corp. warehouse in London on Aug. 9, 2011, the day after it was set on fire by rioters; CDs and record stock belonging to independent record labels were destroyed in the conflagration.


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Date: 3 years ago
Runtime: 0:54
Views: 5,257
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Now THAT'S what I'm talkin bout! Dope. Please do more of these. And longer doggystyle vids too while you're at it. Thanks for the great vids, man.
I find this vid extremely arousing, because of those sexy feet and toes.
dayum dude! u r hott .. ur feet are absolutely immaculate


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Don't Look At Black Holes Too Closely, They Might Disappear
An artist's impression of a supermassive black hole. Black holes are rife with paradoxes, but a new theory suggests that they can all be resolved by agreeing that space and time break down at very small scales. Image credit: ESO/M. Kornmesser
We've come a long way in 13.8 billion years; but despite our impressively extensive understanding of the Universe, there are still a few strings left untied. For one, there is the oft-cited disconnect between general relativity, the physics of the very large, and quantum mechanics, the physics of the very small. Then there is problematic fate of a particle's intrinsic information after it falls into a black hole. Now, a new interpretation of fundamental physics attempts to solve both of these conundrums by making a daring claim: at certain scales, space and time simply do not exist.
Let's start with something that is not in question. Thanks to Einstein's theory of special relativity, we can all agree that the speed of light is constant for all observers. We can also agree that, if you're not a photon, approaching light speed comes with some pretty funky rules – namely, anyone watching you will see your length compress and your watch slow down. [read more...]
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What's That Line? Misheard Lyrics From Some of Music's Biggest Artists
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Page 1
Biophysical properties of Saccharomyces cerevisiae
and their relationship with HOG pathway activation
Jo ¨rg Schaber•Miquel A`ngel Adrover•Emma Eriksson•Serge Pelet•
Elzbieta Petelenz-Kurdziel•Dagmara Klein•Francesc Posas•
Mattias Gokso ¨r•Mathias Peter•Stefan Hohmann•Edda Klipp
Received: 11 January 2010/Revised: 18 March 2010/Accepted: 12 May 2010/Published online: 19 June 2010
? The Author(s) 2010. This article is published with open access at Springerlink.com
mechanical cell properties are important for predictive
mathematical modeling of cellular processes. The concepts
of turgor, cell wall elasticity, osmotically active volume,
and intracellular osmolarity have been investigated for
decades, but a consistent rigorous parameterization of these
concepts is lacking. Here, we subjected several data sets
of minimum volume measurements in yeast obtained
after hyper-osmotic shock to a thermodynamic modeling
framework. We estimated parameters for several relevant
biophysical cell properties and tested alternative hypotheses
Parameterizedmodelsof biophysicaland
In accordance with previous reports, we estimated an aver-
age initial turgor of 0.6 ± 0.2 MPa and found that turgor
becomes negligible at a relative volume of 93.3 ± 6.3%
corresponding to an osmotic shock of 0.4 ± 0.2 Osm/l. At
high stress levels (4 Osm/l), plasmolysis may occur. We
found that the volumetric elastic modulus, a measure of cell
wall elasticity, is 14.3 ± 10.4 MPa. Our model discrimina-
tion analysis suggests that other thermodynamic quantities
affecting the intracellular water potential, for example the
matrix potential, can be neglected under physiological con-
ditions. The parameterized turgor models showed that acti-
vation of the osmosensing high osmolarity glycerol (HOG)
signaling pathway correlates with turgor loss in a 1:1 rela-
tionship. This finding suggests that mechanical properties of
the membrane trigger HOG pathway activation, which can
be represented and quantitatively modeled by turgor.
M. A`. Adrover, E. Eriksson, S. Pelet, E. Petelenz-Kurdziel have
contributed equally to this work.
Electronic supplementary material
article (doi:10.1007/s00249-010-0612-0) contains supplementary
material, which is available to authorized users.
The online version of this
J. Schaber ? E. Klipp (&)
Theoretical Biophysics, Humboldt University,
Invaliden Str. 42, 10115 Berlin, Germany
e-mail: firstname.lastname@example.com
M. A`. Adrover ? F. Posas
Cell Signaling Unit, Departament de Cie `ncies Experimentals i de
la Salut, Universitat Pompeu Fabra (UPF),
08003 Barcelona, Spain
E. Eriksson ? M. Gokso ¨r
Department of Physics, University of Gothenburg,
412 96 Gothenburg, Sweden
S. Pelet ? M. Peter
Institute of Biochemistry, ETH, Zurich, Switzerland
E. Petelenz-Kurdziel ? D. Klein ? S. Hohmann
Department of Cell and Molecular Biology/Microbiology,
University of Gothenburg, Box 462, 405 30 Go ¨teborg, Sweden
Present Address:
D. Klein
Clinical R&D Laboratory, Oxoid, Microbiology,
Basingstoke, Thermo Fisher Scientific, Perth, UK
J. Schaber (&)
Institute for Experimental Internal Medicine, Medical Faculty,
Otto von Guericke University, Leipziger Str. 44,
39120 Magdeburg, Germany
e-mail: firstname.lastname@example.com
Present Address:
E. Eriksson
SP Technical Research Institute of Sweden,
Brinellgatan 4, Box 857, 501 15 Bora ˚s, Sweden
Eur Biophys J (2010) 39:1547–1556
DOI 10.1007/s00249-010-0612-0
Page 2
Volumetric elastic modulus ? High osmolarity glycerol
(HOG) signaling ? Plasmolysis ? Model discrimination ?
Turgor ? Cell wall elasticity ?
All living cells maintain an osmotic pressure gradient
between their interior and the extracellular environment.
This gradient is counterbalanced by a hydrostatic pressure,
called turgor, which is especially prominent in plants and
unicellular organisms, most notably fungi. The concept of
turgor has been debated for many years (Burstro ¨m 1971). It
is recognized that turgor plays a vital role in growth
(Cosgrove 1981; Zimmermann 1978; Marechal and Gervais
1994), cell structure (Cosgrove 1993; Morris et al. 1986;
Zhongcan and Helfrich 1987; Munns et al. 1983), and
membrane transport processes (Zimmermann 1978; Lande
et al. 1995; Kamiya et al. 1963; Soveral et al. 2008) and
that it may be sensed by the cells as an indicator of external
osmotic changes (Tamas et al. 2000; Coster et al. 1976;
Schmalstig and Cosgrove 1988; Reiser et al. 2003).
Therefore, quantifying cell wall and membrane properties,
and thus the resulting turgor pressure, has been the aim of
numerous studies (Cosgrove 1981; Morris et al. 1986;
Munns et al. 1983; Kamiya et al. 1963; Coster et al. 1976;
Smith et al. 1998, 2000a, b, c; Marechal et al. 1995;
Gervais et al. 1996a; Cosgrove 2000). Parameterized
models of turgor pressure, based on cell wall and mem-
brane properties are especially important for quantitative
mathematical models describing the interdependence
between water transport and signaling processes (Schaber
and Klipp 2008; Klipp et al. 2005; Gennemark et al. 2006;
Kargol and Kargol 2003a, 2003).
At present, it is not possible to directly measure turgor
in yeast. However, the so-called linear elastic theory
states that the change in turgor pressure P is proportional
to a relative change in membrane enclosed cell volume
Vm, i.e.,
dP ¼ edVm
where e is a proportionality factor, called volumetric elastic
modulus or bulk modulus. This factor can be viewed as a
measure of the elasticity of the cell wall. Thus, the turgor
pressure can be derived from cell volume measurements
that have been performed repeatedly on yeast cells under
different experimental conditions (Munns et al. 1983;
Smith et al. 2000a, b, c; Touhami et al. 2003; Wei et al.
2001; Martinez de Maranon et al. 1996; Arnold and Lacy
1977). However, the results obtained for turgor pressure
were highly inconsistent, with variations spanning up to
three orders of magnitude from 0.05 (Martinez de Maranon
et al. 1996) to 2.9 (Arnold and Lacy 1977) MPa.
The concepts of turgor, cell wall elasticity, osmotically
active volume and intracellular osmolarity have never been
quantified and parameterized together in a consistent
framework, nor challenged by alternative concepts (Smith
et al. 2000c; Martinez de Maranon et al. 1996; Gervais
et al. 1996b). Moreover, the few reports in which turgor
and/or cell wall elasticity were quantitatively determined
example, one study used pre-stressed cells (Marechal et al.
1995) and in two others, the cells were suspended in dis-
tilled water before volume measurements (Smith et al.
2000b, c).
The objective of this study was to derive parameterized
models of turgor and other biophysical properties of
the cell, for example the volumetric elastic modulus, in
Saccharomyces cerevisiae. We developed a modeling frame-
work that includes the concepts of turgor pressure, intra and
extracellular osmotic pressure, and membrane and volume
properties and, for the first time, also other thermodynamic
quantities, for example the matrix potential, in a consistent
manner. By reformulating sub-modules while keeping
the overall framework consistent, we could test several
hypotheses describing the relationship between cell volume
and changes in the intracellular water potential. Verification
of these hypotheses was based on four independent data
sets. To ensure a wide applicability of the selected model,
the conditions for generating each data set were slightly
different. We also correlated cell volume measurements
with Hog1 phosphorylation levels and Hog1-GFP nuclear
localization to investigate the physiological consequences
of turgor loss under hyper-osmotic stress in a quantitative
fashion. Hog1 is the yeast p38 stress-activated protein
kinase whose phosphorylation level and subcellular locali-
zation are controlled by an osmo-sensing signaling pathway
(Hohmann 2002). We found a 1:1 relationship between
turgor loss and HOG pathway activation. A loss of turgor
relates to HOG pathway activation such that when turgor
becomes zero because of application of a hyperosmotic
solution the HOG pathway is maximally activated. This
finding suggests that the extent of HOG pathway activation
is directly linked to some mechanical property of the
membrane, which can be represented and quantitatively
modeled by the loss of turgor pressure.
conditions. For
Materials and methods
Descriptions and units of all variables and parameters are
listed in Table 1.
1548 Eur Biophys J (2010) 39:1547–1556
Page 3
Modeling framework
We assume that the change in osmotically active volume
Vosis equal to the water flux across the cell membrane,
which is driven by differences in chemical water potentials
inside and outside the cell. Based on these thermodynamic
principles (for details refer to the ''Supplementary mate-
rial'') we developed a model describing the minimal vol-
ume of yeast cells attained after osmotic shock, Vap
included several new aspects that are depicted in Fig. 1.
We made a distinction between the commonly measured
volume that is enclosed by the cell wall, which we refer to
as the apparent volume Vap, and the membrane-enclosed
cytoplasmic volume, Vm, where osmotic and hydrostatic
pressures are defined. Vmcomprises the osmotically active
volume Vosand the minimal solid cytosolic volume Vb, i.e.,
Vm= Vos? Vb. We assume that for osmotic shocks higher
than a critical value of cpl
value of Vap
wall (Vap), i.e., plasmolysis occurs, giving rise to the
min, which
eor volumes lower than a critical
pl, the membrane (Vm) detaches from the cell
periplasmic volume Vpl, such that Vap= Vm? Vpl. Fur-
thermore, we define Vm
becomes zero and Vm
as the volume where negative
hydrostatic effects and/or matrix potential effects become
important, corresponding to hyperosmotic shock levels of
and cs
thresholds that affect the rate of water outflow, i.e.,
shrinkage. Between Vm
reciprocal function of external osmolarity, also called the
van't Hoff relationship (Noble 1969) (Eq. S6 in ''Supple-
mentary material''). Based on these concepts, we derived a
formula for the apparent minimum volume after osmotic
shock Vap
larities and volumes:
aPCRTþ ce
where c0
''Supplementary material''), Vap
P=0as the volume at which the turgor
ee, respectively (Fig. 1). The latter two are
P=0and Vm
sapparent cell volume is a
minas a function of turgor pressure PVmin
ap, osmo-
ap¼ Vbþ Vplþ
ap? Vb? Vpl
stressþ ce
iis the initial internal osmolarity (Eq. S4 in
0is the initial apparent cell
Table 1 Summary of quantities used in the main text
QuantityUnit Description
% initial volume Apparent cell volume, enclosed by the cell wall
% initial volume Initial Vap
Minimal Vapattained after hyperosmotic shock
Cell volume, enclosed by the plasma membrane
% initial volume
% initial volume
% initial volume Osmotically active volume
% initial volume Periplasmic volume. We tested three alternative models of Vplas a function of cstress
''Supplementary material'')
% initial volumeSolid cytosolic volume
% initial volumeVolume, when turgor becomes zero
% initial volumeVolume below which negative hydrostatic or matrix potential effects are important
% initial volumeApparent volume where plasmolysis occurs
Osm/lInitial internal osmolarity.
Osm/l Initial external osmolarity
Osm/l Applied osmotic stress
Initial turgor at the initial turgid cell volume of 100%
corresponding to Vm
corresponding to Vm
corresponding to Vap
e pl
MPa Turgor at Vap
''Supplementary material'') and the two-sided model (Eq. S8 in ''Supplementary material'')
(Fig. 2)
min. In general, there are two alternatives, i.e., the one-sided model (Eq. S7 in
MPaVolumetric elastic modulus, describing the elasticity of the cell wall, relating relative volume
change to hydrostatic pressure, i.e., turgor
MPaProportionality factor, relating relative volume change to the matrix potential or other effects
negatively influencing the intracellular water potential
J K-1mol-1
Gas constant
K Temperature
Conversion factor relating pressure to osmolarity, e.g., for relating MPa and Osm/l it is 10-3
Eur Biophys J (2010) 39:1547–1556 1549
Page 4
volume, R is the gas constant, T is the temperature, aPCis a
dimensionless conversion factor, c0
osmolarity, and cstress
is the applied osmotic stress.
eis the initial external
Turgor pressure and matrix potential
Thermodynamic quantities other than turgor and osmolar-
ity, that can affect the cellular water potential, are usually
ignored. For example, a specific fraction of the intracellular
water might be bound to colloidal surfaces by electrostatic
interactions. This water is not freely available in solution,
which lowers the total intracellular water potential by a
value that is termed the matrix potential (Griffin 1981). In
addition, it is possible that the cytoskeleton mechanically
counteracts cell shrinkage such that the cell has an inherent
tendency to re-swell in a sponge-like fashion. This effect
would also reduce the total intracellular water potential.
For simplicity, we summarize these effects under the term
matrix potential (Ds in Eq. S1 of the ''Supplementary
material''). Quantitative descriptions of the matrix poten-
tial are not available and elaborated models of mechanical
forces induced by volume changes (Stamenovic and
Coughlin 1999) are beyond the scope of this study.
Therefore, we include the effects summarized under the
term of the matrix potential in the model of the turgor
pressure PVmin
with a simple mathematical description. In
this way, we obtain an expression for PVmin
matrix potential (assuming Vpl= 0 for VmC Vm
0 for
including the
ap? Vpl\VP¼0
ap? Vpl\Vs
InFig. 2
min- Vpl.
We tested Eq. 3 against the data, with and without the
term for matrix potential, which we refer to as the one-
sided and the two-sided models, respectively. We also
tested three hypotheses regarding the size of the periplas-
mic volume Vpl, (a) Vpl= 0, (b) Vpl[0 with a constant
apparent volume Vap
material''), and (c) Vpl[0 with a variable apparent volume
Inserting Eq. 3 into Eq. 2 yields an implicit expression
for Vap
model for turgor, matrix effect, and periplasmic volume.
When a solution exists, this solution is unique (see ''Sup-
plementary material'').
isdisplayed asafunction
Vm= Vap
pl(Fig. 1; Eq. S10 in ''Supplementary
pl(Eq. S11 in ''Supplementary material'').
minthat depends on various parameters of the specific
Fig. 1 Qualitative representation of cell volume concepts. Solid line:
Vap, the apparent cell wall enclosed volume. Dotted line: Vm, the
membrane enclosed volume. Vm
becomes zero and Vm
is the volume where negative hydrostatic
effects or matrix potential effects become important, corresponding to
hyperosmotic shock levels of cP=0
and cs
the stress and volume thresholds, respectively, where Vmdetaches
from Vap. Vplis the periplasmic volume. Vbis the minimum solid
cytosolic volume. Note, that it is not necessarily true that cs
membrane in different states after hyper-osmotic shock
P=0is the volume at which the turgor
ee, respectively. cpl
eand Vap
pl. The diagrams of the cells illustrate the cell wall and the
Fig. 2 Illustration of turgor and matrix potential. Turgor (black line),
which has an initial value of P0at the initial turgid cell volume of
100%, is displayed on the positive y-axis. When the cell shrinks, the
turgor decreases to zero at the volume Vm
''one-sided turgor model''. The matrix potential or other effects that
reduce the water potential are displayed on the negative y-axis. These
are assumed to become important below a specific threshold volume
the dashed line is referred to as the ''two-sided turgor model''
P=0. This is referred to as the
s. The model represented by the solid line up to Vm
stogether with
1550 Eur Biophys J (2010) 39:1547–1556
Page 5
Model discrimination
Using Eq. 2 together with a model for Vpl(Eqs. S10, S11 in
''Supplementary material'') and a turgor model (Eq. 3;
Eqs. S7, S8 in ''Supplementary material''), a solution for
sured, Eq. 2 can be fitted to the data by estimating the
respective parameters (Table S1 in ''Supplementary mate-
rial''), e.g., by minimizing the sum of squared residuals
where x = (x1, x2, …, xm) denote the parameters that are
to be estimated and that depend on the model (Table S1 in
''Supplementary material''), riis the standard deviation of
the measurements and n is the number of data points. For
the sake of simplicity, we assume that the measurement
errors are independently normally distributed. Thus, the
SSR becomes a maximum-likelihood estimator and we can
use the Akaike's information criterion corrected for small
sample size (AICc) to rank and, thus, discriminate the
models according to their SSR and number of parameters
(Burnham and Anderson 2002):
where k is the number of parameters.
The fitting was performed by differential evolution, a
global optimization method (Storn and Price 1997), and
the models were ranked according to the AICc (Eq. 4).
Additionally, for the best model, we performed a Monte–
Carlo analysis by re-sampling the data a hundred times
assuming a normal distribution with the respective stan-
dard deviations for each data point. In this way, means
and confidence intervals for the estimated parameters
could be obtained.
mincan be numerically determined. When Vap
minis mea-
SSRðxÞ ¼
ap;iðxÞ ? Vmin
AICc ¼ 2k þ n ln
þ 1
þ2kðk þ 1Þ
n ? k ? 1;
Data sets
We collected three independent data sets (data sets 1–3 in
Fig. 3; Fig. S1 in ''Supplementary material'') and took one
data set from the literature (Marechal et al. 1995) (data set
4 in Fig. 3; Fig. S2 in ''Supplementary material''). The cell
volume was quantified immediately before and after the
hyperosmotic stress, ensuring that the minimum volume
after osmotic shock was measured. For our experiments we
used NaCl as an osmoticum, whereas in the data set from
literature yeast cells were subjected to a wide range
of glycerol concentrations (refer to Marechal et al. 1995
for details). Table 2 summarizes the different data sets.
Experimental details for data sets 1–3 are described in the
''Supplementary material''.
Parameter fitting and model selection predicts turgor
pressure and other biophysical cell properties
We fitted Vap
alternative models for PVmin
different models (see Table S1 in ''Supplementary mate-
rial'' for a complete listing). For each data set the best
model according to the AICc is described in Table 3. The
fitted and measured Vap
corresponding predicted turgor pressure.
minfrom Eq. 2 to four different data sets using
and Vpl, resulting in seven
minare displayed in Fig. 3, with the
One-sided turgor model is most appropriate
for physiological stress conditions
The three data sets produced for this study were all fitted
best by the one-sided turgor model based on Eqs. 1 and 3,
where only the positive turgor pressure plays a role
(Fig. 2). Even though this is the generally accepted model
(Cosgrove 1981; Marechal and Gervais 1994; Klipp et al.
2005; Gennemark et al. 2006; Meikle et al. 1988), we
tested it with six other models, because to our knowledge it
has never been challenged by other models, nor rigorously
fitted to data. Thus, we show for the first time that effects
resisting cell shrinkage, for example the matrix potential or
mechanical forces, can be neglected under physiological
conditions. Consequently, when the turgor becomes zero
after an external osmotic shock of cP=0
the measured apparent volume Vapis well predicted by the
membrane enclosed volume Vm according to the van't
Hoff's equation (Eq. S6 in ''Supplementary material'').
This means that the cell wall tightly follows the shrinkage
of the membrane enclosed volume. Data set 4, which was
collected by applying extreme stress levels, also favored
the one-sided turgor model. However, at 4 Osm/l plas-
molysis was predicted, with a constant minimal apparent
volume Vap
(Table 1; Fig. 3),
plof 65%.
Estimated biophysical cell properties are similar
between experiments
Parameters estimated on the basis of the datasets give a
consistent picture of important biophysical cell properties.
For data sets 2 and 3, where single cell information was
available, we distinguished between small and large cells
and repeated the analysis to test whether cell size had an
effect on the results. We found no significant differences
between small and large cells with regard to the estimated
parameters, which is in agreement with earlier reports
(Martinez de Maranon et al. 1996). The solid volume Vb
was estimated to be between 33 and 49% of the initial
cell volume, which also corresponds to earlier reports
Eur Biophys J (2010) 39:1547–1556 1551
Page 6
(Marechal et al. 1995; Meikle et al. 1988). The initial
turgor estimated on the basis of the three data sets with
similar experimental conditions (data sets 1–3) was
0.6 ± 0.2 MPa on average, which confirmed results from
earlier studies (Smith et al. 2000c; Gervais et al. 1996a;
Meikle et al. 1988). The estimated membrane rigidity e was
the parameter that varied the most between experiments
and it turned out to be a sensitive, poorly determined
parameter, which is reflected by the relatively high stan-
dard deviations. However, at least for the three directly
comparable data sets 1–3, the results were similar, giving
an average volumetric elastic modulus of 14.3 ± 10.4 MPa
ranging from 5.7 to 25.8 MPa.
On the basis of our general modeling framework and the
estimated parameters, we could derive several other inter-
esting cell properties. The initial osmolarity within the cell
of the outside medium. Thus, apparently, yeast can main-
tain substantial osmotic gradients between the inside and
outside of the cell, which are equilibrated by turgor. The
volume at which turgor becomes zero was estimated to be
of 93.3 ± 6.3% on average for data sets 1–3. The corre-
sponding external shock for the salt experiments was
0.4 ± 0.2 Osm/l, corresponding to a salt concentration of
roughly 0.2 M NaCl. As this is within the range of con-
centrations in which saturation of HOG pathway activation
iwas estimated to be up to more than twice the osmolarity
Fig. 3 Best fit models and data. The data and the best fit model (solid
line) refer to the left y-axis. The dark gray region depicts the 95%
confidence region of the fitted model. The right y-axis refers to
the estimated turgor pressure (dashed line). The light gray region
depicts the 95% confidence region of the estimated turgor pressure.
The x-axis refers to concentrations of the respective stress agent. The
confidence regions were calculated on the basis of the confidence
intervals of the estimated parameters (Table 3). The four data sets
are described in the ''Methods'' sections and the ''Supplementary
Table 2 Data sets used for model discrimination
No. MethodStrainMedium Agent
e(Osm/l)# n
1Particle sizerW303 YPD NaCl30Early log0.26 16 This study
2 Single cells bright fieldBY4741 YNBNaCl20 Log 0.2711 This study
3 Single cells fluorescenceW303 SDNaCl 30Log 0.25 13 This study
4Single cells bright field CBS1171 WickerhamGlycerol 25 Stationary0.86 15Marechal et al. (1995)
ewas measured for data sets 1–3 with an osmometer and calculated for data set 4 from Marechal et al. (1995), # is the number of data points
1552Eur Biophys J (2010) 39:1547–1556
Page 7
is observed (Maeda et al. 1995), we hypothesized that the
initial HOG pathway response may be correlated with
Loss of turgor correlates with HOG pathway activation
The exact biophysical mechanisms that trigger the activa-
tion of the HOG pathway in yeast have not yet been
determined (Hohmann 2002), but there is evidence that the
turgor is one of the factors involved (Tamas et al. 2000;
Reiser et al. 2003). As our computational analysis provided
us with parameterized models of turgor pressure (Fig. 3;
Table 3), we tested whether there is a correlation between
the loss of turgor for different degrees of osmotic shock
and the extent of HOG pathway activation. Thus, in addi-
tion to cell volume measurements, we collected two inde-
pendent data sets reflecting the degree of HOG pathway
activation depending on the intensity of the hyperosmotic
stress. In the first data set, the extent of Hog1 nuclear
accumulation was measured, whereas the second data set
represents the extent of Hog1 phosphorylation in response
to different NaCl concentrations (Fig. 4, for details about
the measurements refer to the ''Supplementary material'').
By considering both Hog1 activation (Fig. 4) and the
predicted turgor (dotted line in Fig. 3) as a function of
applied osmotic stress, we could investigate Hog1 activa-
tion as a function of predicted turgor (Fig. 5). With three
different parameterizations for the turgor model (from data
sets 1–3, data set 4 was disregarded, because it used a
different stress agent) and two different Hog1 activation
data sets, we obtained six turgor-Hog1 activation correla-
tions, of which we display three in Fig. 5. The others are
displayed in the ''Supplementary material'' (Fig. S3).
We fitted a linear relationship between HOG pathway
activation and relative turgor for the three different
parameterized models, corresponding to data sets 1–3, and
the two different HOG pathway activation data sets (black
lines in Fig. 5) by a weighted orthogonal regression. Our
null hypothesis claimed a direct 1:1 linear relationship,
i.e., H0: a ? bx, with (a, b) = (100, -1) (gray lines in
Fig. 5). To test H0we computed confidence regions for
the estimated parameter pairs (a, b) by a Monte–Carlo
analysis. As can be seen from the insets in Fig. 5, in all
cases the hypothesized relationship (gray points) was
within the 95% confidence region of the actual estimated
parameter pair. Thus, the null hypothesis could not be
Table 3 Results of parameter fits of Eq. 2 for the best selected model according to the AICc
Data set 1Data set 2 Data set 3 Data set 4
#2224 Selected model
One-sided One-sided One-sidedOne-sided
000 Constant Vap
49.0 ± 3.4 40.2 ± 5.533.0 ± 4.8 40.1 ± 11.1 Estimated parameters
0.5 ± 0.1 0.9 ± 0.2 0.5 ± 0.13.4 ± 1.0
25.8 ± 7.0 5.7 ± 2.011.4 ± 2.128.6 ± 11.3
65.0 ± 5.9
0.4 ± 0.10.6 ± 0.10.5 ± 0.0 2.2 ± 0.4 Derived parameters
98.2 ± 3.1 86.1 ± 6.395.5 ± 6.8 88.9 ± 10.8
0.2 ± 0.10.5 ± 0.2 0.2 ± 0.11.9 ± 1.9
4.4 ± 2.2
For a complete description of parameters and units refer to Table 1
# Number of best selected model (Table S1 in ''Supplementary material''), PVmin
(Fig. 1; Eq. S10 in ''Supplementary material''), k: number of fitted parameters, Vb: solid base volume, P0: initial turgor, e: volumetric elastic
modulus, Vap
to equilibrate turgor, cpl
the data points were sampled from their respective standard deviations indicated in Figs. 3 and 4. The mean of parameters in the lower part are
derived from the mean parameters in the upper part
m: selected turgor model (Fig. 2; Eq. 3), Vpl: selected Vplmodel
plapparent volume where plasmolysis occurs, c0
e: applied external stress to reach Vap
i: initial internal osmolarity, Vm
pl. Estimated parameters are given as mean ± standard deviation from 100 fits where
P=0: volume at zero turgor, cP=0
: applied external stress
Fig. 4 Hog1 activation data. Circles: maximum Hog1 nuclear
concentration after osmotic shock. Squares: Hog1 phosphorylation
after 2 min of shock treatment. Data are scaled to the respective
measured maximum
Eur Biophys J (2010) 39:1547–15561553
Page 8
rejected. Moreover, the Kendall-rank correlation coeffi-
cient (Kendall and Gibbons 1990) was highly significant
in all cases (P\0.01).
Parameterized models of biophysical and mechanical cell
properties become increasingly important for quantitative
and predictive descriptions of cellular behavior. Our
modeling framework, which included the concepts of tur-
gor, membrane elasticity, intracellular osmolarity, osmoti-
cally active volume, non-turgid volume, and solid cytosolic
volume in a coherent and thermodynamically accurate
way, enabled us to parameterize all these concepts con-
sistently, by fitting the relevant parameters to experimental
data. In addition, it enabled us to discriminate between
alternative hypotheses for turgor and volume changes
by including alternative sub-models within the general
framework. Even though the data sets were based on very
different techniques, the overall conclusions are strikingly
similar indicatingthatthe
description is both accurate and universal.
We confirmed previous reports, which suggested an
initial turgor pressure of around 0.6 MPa (Smith et al.
2000b; Meikle et al. 1988). Turgor pressures reported for
yeast differ substantially. Part of this variation might be
because they have never been rigorously fitted to data. In
our study, we obtained similar results for data sets 1–3.
However, the initial turgor estimated from data set 4 was
sixfold higher. We assume that this discrepancy is caused
by different growth phases of yeast cells employed in the
experiments. This explanation is supported by earlier
reports, in which differences in turgor between cells in
different physiological states were observed. Specifically,
a higher turgor pressure was reported for cells in sta-
tionary phase (Smith et al. 2000c; Martinez de Maranon
et al. 1996). After examining different possible explana-
tions (Eq. 3), we confirmed the hypothesis that turgor
pressure is significant only above a specific volume
threshold, below which it can be neglected, as previously
proposed (Cosgrove 1981; Marechal and Gervais 1994;
Klipp et al. 2005; Gennemark et al. 2006; Meikle et al.
1988). Other forces potentially affecting the water
potential, and therefore water flux within the cell, can be
neglected under the studied conditions of osmotic shock
up to 1.5 M NaCl.
We found values for the volumetric elastic modulus e
between 6 and 29 MPa. We could not find any published
values of e for yeast. In a recent study, where yeast cells
were subjected to mechanical compression, values of
Fig. 5 Turgor-HOG pathway activation relationship. The x-axes
are relative turgor (%) as predicted by the parameterized models
for different shocks of NaCl (dotted line in Fig. 3). The y-axes are
relative HOG pathway activation (%) according to different shocks
of NaCl (Fig. 4). The black lines are a fitted linear relationship
(y = a ? bx) based on a weighted orthogonal regression. The gray
lines represent the null hypothesis H0: y = 100 - x, i.e., a direct
1:1 linear relationship between relative loss of turgor and relative
HOG pathway activation. The insets are plots of the (25, 50, 75,
90, 95%)-confidence regions of the respective estimated parameter
pair (a, b) of a ? bx with the outermost line being the 95%
confidence region. The black points correspond to the black lines
in the plots, the gray points correspond to the gray lines. The
confidence regions are obtained by a Monte–Carlo analysis with
1,000 runs
1554Eur Biophys J (2010) 39:1547–1556
Page 9
Young's modulus E for the cell wall of around 110 MPa
were reported (Smith et al. 2000a, c). Unfortunately, we
could not relate these values to our volumetric elastic
modulus, because those values for Young's modulus E
were derived assuming a Poisson's ratio m of 0.5 and,
hence, the formula E = 3e (1 - 2m) was not applicable.
We noted that cells in stationary phase (data set 4) are
estimated to be more rigid, i.e., have a higher volumetric
elastic modulus, but to a much lesser extent than in Smith
et al. (2000c). The cells used in data set 1 were grown to
saturation, i.e., beyond proliferation, before they were re-
suspended in fresh medium for 1 h before measurements.
It is possible that they were still recovering from satura-
tion phase and therefore had higher cell wall rigidity than
cells from data sets 2 and 3, which were measured at
logarithmic growth phase. This reflects that e is a function of
the physiological state of the cell. Thus, when turgor is being
modeled over longer time intervals than in this study, the
assumption of e being constant might be compromised.
Our modeling results support the notion of a rather
elastic cell wall, because the cell wall tightly follows the
cell membrane on shrinkage, and plasmolysis only occurs
under extremely high stresses. There is experimental evi-
dence from mammalian cells that our model of an elastic
membrane/cell wall surrounding a viscous medium is an
over-simplification (Wang et al. 2001). However, our
model proved sufficient to explain our data. More com-
plicated models were not supported by the data in respect
of the AICc. It is known that upon osmotic shock the actin
cytoskeleton in yeast de-polymerizes (Brewster and Gustin
1994). This is in line with our results in which effects of the
skeleton, which we included into the model of the matrix
potential, were refuted by the model discrimination. A
recent compression study of mammalian cells also ruled
out the cytoskeleton as being responsible for the increase in
cell stiffness as the volume decreases (Zhou et al. 2009).
Other authors identified the concentration at which
plasmolysis occurs as approximately 1 Osm/l (Arnold and
Lacy 1977), but under different experimental conditions.
The predicted plasmolysis point of 65% in data set 4 was
higher than the lowest volumes measured in the other data
sets (44–54%, Fig. 3). Therefore, cells might have been at
the threshold of plasmolysis in those experiments. Because
no data were collected at higher stress levels this effect was
not distinguishable by model selection.
For the first time, we provide evidence that there is
direct link between loss of turgor upon osmotic shock and
HOG pathway activation, not only qualitatively but also
quantitatively. Moreover, the hypothesis of a direct 1:1
relationship between turgor loss and HOG pathway acti-
vation could not be rejected. The physical basis of such a
direct relationship remains elusive, but could for example
be a change in Sln1 and Sho1 membrane protein
conformation as a function of membrane stretch or other
mechanical forces that relate to turgor.
ital version of data set 4, Rosie Perkins, Javier Macia, and Karlheinz
Schaber for useful suggestions on the manuscript, and Martina
Fro ¨hlich for measurement support. This work was supported via
several projects funded by the European Commission: QUASI
(Contract No. 503230 to SH, EK, FP and MP), CELLCOMPUT
(Contract No. 043310 to SH, EK and FP), UNICELLSYS (Contract
No. 201142 to SH, EK, FP, MP and MG), SYSTEMSBIOLOGY
(Contract No. 514169 to SH and EK), and AMPKIN (Contract No.
518181 to SH and MG). In addition work was funded by grants from
the Swedish Foundation for Strategic Research SSF (Bio-X to MG),
the Swedish Research Council (project grants to SH and MG), the
Carl Trygger Foundation (to MG), the Science Faculty, University of
Gothenburg (to SH and MG), and the Swiss systemsX.ch (to MP).
We thank Patrick Gervais for providing a dig-
Open Access
Creative Commons Attribution Noncommercial License which per-
mits any noncommercial use, distribution, and reproduction in any
medium, provided the original author(s) and source are credited.
This article is distributed under the terms of the
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1556Eur Biophys J (2010) 39:1547–1556


text:
Music News Archives - Page 142 of 952 - THE WOLF Sacramento
Miranda Lambert's Redemption Ranch Opens in Oklahoma
Miranda Lambert has once again stepped up to help animals in need. Her privately-funded no-kill animal shelter, Redemption Ranch, opened over the weekend in her hometown of Tishomingo, Okla. Continue… MORE
Taylor Swift Exhibit to Open at the Grammy Museum
The Grammy Museum in Los Angeles will soon open 'The Taylor Swift Experience,' an exhibit dedicated to the formerly country music icon. Continue reading…
Top 10 'The Voice' Contestants Join Together for Rascal Flatts Mashup [Watch]
The Top 10 contestants on 'The Voice' came together at the beginning of Monday night's (Nov. 24) show for an incredible Rascal Flatts medley. Continue reading…
Reba McEntire Asks Fans to 'Pray for Peace,' Garth Brooks Cancels Press in Light of Ferguson
Reba McEntire is calling on fans to pray for peace as violence has once again erupted in Ferguson, Mo. Continue reading…
Jake Owen Says Poignant 'What We Ain't Got' Video Portrays the Song's Message
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Dakota Bradley Opens Up About the Misfortune That Brought Him to Nashville, Tim McGraw's Impact and More
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text:
3137 Kerry Court
Dubuque, Iowa, 52001
Commute to Downtown Dubuque
Almost all errands require a car.
Score Details
3137 Kerry Court has a Walk Score of 13 out of 100. This location is a Car-Dependent neighborhood so almost all errands require a car.
This location is in Dubuque. Nearby parks include Veterans Memorial Park, Dubuque Arboretum and Botanical Gardens and Marshall Park.
Nearby schools include Eisenhower Elementary School, Mazzuchelli Catholic Middle School and Wahlert Catholic High School.
The closest coffee shop is One Mean Bean. Nearby restaurants include Dubest Pizza Inc, Godfather's Pizza and Pickle-Barrell Subs.
3137 Kerry Court is near Emmaus Bible College, Gay Park and Clarke University.
Explore how far you can travel by car, bus, bike and foot from 3137 Kerry Court.
View all Dubuque apartments on a map.
This location is in the city of Dubuque, IA. Dubuque has an average Walk Score of 40 and has 57,637 residents.


text:
Weekend Magazine: Senior Depression, Mississippi Writer, And Free Clothing
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College Football Spotlight: College Football's Best Month Will Get Started With A Bang
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At 88, The King of The Blues Is Still Rocking Audiences
Blues guitar legend B.B. King — who played with Eric Clapton and U2 — has been performing for several decades and shows no signs of stopping…
San Jose Solar Firm Goes Under; Mississippi Among Creditors Owed Millions
Mississippi taxpayers may have only an empty Senatobia building and solar panel equipment to show for $26 million in loans to Twin Creeks Technologies.


text:
Toyota has issued a worldwide recall for about 437,000 of its flagship Prius and other hybrid vehicles in the latest of a series of embarrassing safety problems to hit the Japanese carmaker.
"We have decided to recall as we regard safety for our customers as our foremost priority," Akio Toyoda, the company's president, told a press conference in Tokyo on Tuesday.
The recall follows complaints from Toyota owners in the US and Japan about a delay under some condition when the brakes in the Prius are pressed.
Toyota officials had earlier informed Japan's transport ministry about the Prius recall, with grim-faced company bosses bowing before ranks of assembled media.
Toyota said it was also recalling the Lexus HS250h luxury hybrid sedan, the Sai hybrid sold only in Japan and the Prius plug-in hybrid which is an experimental model used on a small scale in Japan.
Software glitch
The recall in Japan covers about 223,000 vehicles, with most of the rest of the affected cars thought to have been sold in the US – Toyota's biggest market.
|Toyota Prius
World's first mass-produced gas-electric hybrid car
Launched in Japan in 1997, now sold in more than 40 countries
Third generation model launched in May 2009
Listed by US Environmental Protection Agency as the most fuel efficient car on the market
More than 1.6 million Prius cars sold around the world, half of them in the US
World's best-selling hybrid vehicle that has helped Toyota grab 70 per cent of the US market for such vehicles
Toyota says it has fixed the software programming glitch in Prius vehicles that have gone on sale since last month, but not those sold before, starting last May.
"Let me reassure that we will redouble our commitment to quality as the lifeline of our company," Toyoda told reporters at Tuesday's press conference.
The company's reputation for quality and reliability has been severely dented in recent weeks by a series of recalls of around eight million vehicles due to sticking accelerator pedals.
Vignesh Chandran, an auto industry specialist in London, told Al Jazeera: "If you look at real safety issues, there are regulatory boards that test cars before they go to market.
"It is consumer confidence that has really been affected. It is too early for us to say these cars are not safe."
Toyota says it has fixed the programming glitch in Prius vehicles that have gone on sale since last month, but not those sold before, starting last May.
The Japanese media and politicians have been critical of its slow response and lack of action plan, and drivers told our reporter they felt betrayed by the company.
Toyota had been trying to make out the Prius braking issue as a minor quirk, but in the wake of its recent safety recalls with other models, many consumers feel that is just not good enough, he said.
The world's top car maker has already recalled more than 8 million other vehicles worldwide in recent weeks because of safety concerns centred on floor mats and sticky accelerator pedals that can cause unintended acceleration.
The company's North American plants have resumed production of eight models involved in the earlier safety recalls now but faces a potential rush of litigation.
At least 30 class-action lawsuits have been filed against the Japanese car maker over the accelerator pedals and on Friday it faced its first US suit over complaints of faulty braking in its Prius model.
The National Highway Traffic Safety Administration (NHTSA) in the US says it has opened a formal investigation into Prius-related complaints of "momentary loss of braking capability while travelling over an uneven road surface, pothole or bump".
The NHTSA has received nearly 200 such complaints since February 3, including four reports of accidents, two of them said to have resulted in injury.
Despite its problems, Toyota's shares, which have lost about a fifth of their value since late January, were up 1.7 per cent early on Tuesday.
Hiroaki Osakabe, a fund manager at Chibagin Asset Management, said Toyota shares fell while the company "appeared not to be doing anything to deal with its problems".
"But now, the fact that they're taking concrete steps on the issue is being seen as positive," he said.
He warned, however, that "the gains will be limited since many investors want to see the impact of the problems on US car sales, with more share selling ahead if there's a big drop".
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Xcel billing issues in North Dakota spotlight
BISMARCK - A newly-elected North Dakota Public Service Commission member is taking a serious look at billing or meter problems occurring at the state's largest utility.
Public Service Commissioner Brian Kalk, who took office Jan. 1, is asking questions about Xcel Energy's use of estimated bills for multiple months this winter, which have resulted in electric heat bills this month of up to $900 for some residential customers and similar jolts for a few natural gas customers.
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Other titles in the Que Biz-Tech series:
Pinterest for Business: How to Pin Your Company to the Top of the Hottest Social Media Network (Que Biz-Tech)by Jess Loren
Synopses & Reviews
Publisher Comments:
The complete guide to profiting from Pinterest!
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About the Author
Jess Loren is a consultant, strategist, speaker, and eternal "lemonade stand" purveyor (aka, entrepreneur) in the world of social media and digital marketing. She is the cofounder and managing partner of Kambio Group, a digital marketing agency that concentrates on social media management and event-based campaigns. Loren has contributed across the industry and across the spectrum, working with everyone from Fortune 500 companies to a company that hosts the "largest bar crawl in America."
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Table of Contents
1. Welcome to Pinterest: An Informal Introduction:
2. The Nitty Gritty: Pinterest in Your Businesses:
3. Pinpointing Your Target: Captivating Your Audience
4. Pinpointing Your Industry: Who Am I, and Who Are They
5. Winning by Pinning Within Pinterest Categories: Home/Crafts/Beauty/Entertainment
6. Winning by Pinning Within Pinterest Categories: Food/Apparel/Shoes/Travel/Celebrity
7. Relentless Reach: the Power Behind Your Pins
8. Viralocity: How to Improve Your Pin's Reach
9. Playing Well With Others: Pinterest and Other Social Media Brands
10. Being the Bright Pin: Standing Out On a Crowded Pinboard
11. The Hidden Rules of Pinterest: Do's and Don'ts
12. Conclusion
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Learn more about the five boroughs that make up New York City with our complete guide.
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NYC is the place for inquisitive minds looking for new things to see, hear, taste and learn.
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NYC Comedy Week – 2-for-1 Tickets to New York Comedy Shows
March 23–29, celebrate the show's final season with a two-course lunch or two cocktails for $19.69.
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New York City is made up of five boroughs: the Bronx, Brooklyn, Manhattan, Queens and Staten Island. Each one has enough attractions—and enough personality—to be a city all its own. Learn more about them with this guide.
Established in 1864, Pete's is a contender for the City's oldest continuously operating bar. Even Prohibition couldn't prevent the place from closing its doors—it transformed into a speakeasy with a flower shop as its front. Lore has it that O. Henry wrote his 1906 short story "The Gift of the Magi" at one of the front tables. Today, patrons enjoy its burgers and old-school Italian food along with the house brew, Pete's 1864 Original House Ale.