text:
Strip steak
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Strip steak
Beef Cuts
Type
Short loin cut of beef
Grilled Kansas City strip steak topped with onions and mushrooms
The strip steak is a cut of beef steaks from the short loin from a cow. It consists of a muscle that does little work, the longissimus, making the meat particularly tender;[1] although not as tender as the nearby psoas major or tenderloin. Fat content of the strip is somewhere between the two cuts. Unlike the tenderloin, the longissimus is a sizable muscle, allowing it to be cut into larger portions.
Contents
[hide]
1 Other names
2 Related cuts
3 See also
4 References
Other names[edit]
A U.S. Prime raw strip steak with a high marbling content
Raw strip steak
According to the National Cattlemen's Beef Association, the steak is marketed in the United States under various names, including Ambassador Steak, Boneless Club Steak, Hotel-Style Steak, Kansas City Steak, New York Steak, Top Loin, and Veiny Steak.[2]
In New Zealand and Australia, it is known as Porterhouse and Sirloin (striploin steak)[3] and can be found in the Handbook of Australian Meat under codes 2140 to 2143.[4]
In the UK and Ireland it is called sirloin.
In Canada, most meat purveyors refer to this cut as a strip loin;[5] in French it is known as contre-filet.
Delmonico's Restaurant, an operation opened in New York City in 1827, offered as one of its signature dishes a cut from the short loin called a Delmonico steak. Due to its association with the city, it is often referred to as a New York strip steak.[6]
Related cuts[edit]
When still attached to the bone, and with a piece of the tenderloin also included, the strip steak becomes a T-bone steak or a porterhouse steak, the difference being that the porterhouse has a larger portion of tenderloin included. The strip steak may be sold with or without the bone. Strip steaks may be substituted for most recipes calling for T-bone and porterhouse steaks, and sometimes for fillet and rib eye steaks.
A bone-in strip steak with no tenderloin attached is sometimes referred to as a shell steak.[7]
See also[edit]
Food portal
British and American cuts of beef
Entrecôte
References[edit]
Jump up ^ Herbst, Sharon. "Kansas City Strip Steak". Epicurious. Barron's Educational Services. Retrieved 28 November 2011.
Jump up ^ "Understanding the Cuts". farmfreshbeef.org. Retrieved 2014-10-30.
Jump up ^ "Beef Cuts Chart" (PDF). australian-beef.com. Meat & Livestock Australia, Ltd. Retrieved 2016-11-13.
Jump up ^ "Australian RFP Cut Code Reference 2016 Edition" (PDF). ausmeat.com.au. Aus-Meat, Ltd. Retrieved 2016-11-13.
Jump up ^ "Beef - Meat Cuts Manual". inspection.gc.ca. Canadian Food Inspection Agency. Retrieved 2016-04-07.
Jump up ^ goodtastebook.com
Jump up ^ http://www.seriouseats.com/2011/03/the-four-high-end-steaks-you-should-know-ribeye-strip-tenderloin-t-bone.html
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Related meats
American bison
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Bovine spongiform encephalopathy
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Carcass grade
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text:
I Just Don't Know What to Do with Myself
From Wikipedia, the free encyclopedia
Jump to: navigation, search
This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (March 2016) (Learn how and when to remove this template message)
"I Just Don't Know What to Do with Myself"
Single by Dusty Springfield
from the album Dusty
B-side
"My Colouring Book"
Released
1964
Format
7"
Genre
Pop
Length
3:01
Label
Philips
Songwriter(s)
Burt Bacharach and Hal David
Producer(s)
Johnny Franz
Dusty Springfield singles chronology
"All Cried Out"
(1964) "I Just Don't Know What to Do with Myself"
(1964) "Guess Who"
(1964)
"All Cried Out"
(1964) "I Just Don't Know What to Do with Myself"
(1964) "Guess Who"
(1964)
"I Just Don't Know What to Do with Myself" is a song written by Burt Bacharach and Hal David.
Contents
[hide]
1 Original version
2 Dusty Springfield version
2.1 Charts
3 Dionne Warwick version
3.1 Formats and track listing
3.2 Charts
4 Marcia Hines version
4.1 Track listing
4.2 Weekly charts
4.3 Year-end charts
4.4 See also
5 White Stripes formats and track listing
6 Other versions
7 Bibliography
8 References
9 External links
Original version[edit]
"I Just Don't Know What to Do With Myself" was first recorded by Tommy Hunt in a session produced by Jerry Leiber and Mike Stoller, with Burt Bacharach arranging and conducting. It was released as single Scepter 1236 (B-side "And I Never Knew") in May 1962, but did not chart. The song served as the title track for Hunt's sole album release on Scepter released in April 1963. The success in the UK of the Dusty Springfield version of "I Just Don't Know..." in the summer of 1964 led to Scepter's re-releasing the original, at which time it reached the Bubbling Under Hot 100 Singles chart in Billboard with a #119 peak. It would appear however that the first version of this song to be recorded was not by Hunt but by Chuck Jackson also in 1962. Jackson's version was shelved and remained unreleased until it appeared on a 1984 compilation on the Kent record label called "Mr. Emotion". According to the sleeve notes of that album, Tommy Hunt's vocals were substituted for Jackson's whilst the original backing track was retained.
Dusty Springfield version[edit]
Dusty Springfield recorded "I Just Don't Know What to Do with Myself" in a session at Olympic Studios in Westminster with production credited to Philips Records owner Johnny Franz - although Springfield later stated her solo Philips tracks were self-produced - and arranged by Ivor Raymonde who conducted his orchestra; personnel on the session included Big Jim Sullivan on guitar and Bobby Graham on drums. Springfield, whose first solo recordings had included covers of the Bacharach/David compositions "Anyone Who Had a Heart" and "Wishin' and Hopin' " - had brought back "I Just Don't Know..." from an overnight trip to New York City where she met up with Bacharach in February 1964.
The third UK single release of Springfield's solo career - following the "Brill Building" Sound-alikes "I Only Want to Be With You" and "Stay Awhile" - "I Just Don't Know What to Do with Myself" was Springfield's first UK single release to display her signature vocal style; rising to #3 in the summer of 1964 the track remained Springfield's highest charting UK hit until she reached #1 in 1966 with "You Don't Have to Say You Love Me" which would remain Springfield's only UK solo hit to chart higher than "I Just Don't Know...".
A concurrent US release of Springfield's "I Just Don't Know What to Do with Myself" was preempted by the presence of Springfield's "Wishin' and Hopin'" in the US Top Ten over the summer of 1964. Springfield's "I Just Don't Know..." received a belated US release in October 1965 featured on a single with Springfield's current UK hit "Some of Your Lovin'"; that month Springfield made a promotional junket to the US which included performances of both of the single's tracks on the TV shows Hullabaloo and Shindig but neither side reached the Billboard Hot 100.
Charts[edit]
Chart (1964)
Peak
position
UK Singles Chart[1] 3
Australian Singles Chart[2] 16
Dutch Singles Chart[3] 5
Dionne Warwick version[edit]
"I Just Don't Know What to Do with Myself"
Single by Dionne Warwick
from the album Here Where There Is Love
B-side
"In Between the Heartaches"
Released
1966
Format
7" Vinyl
Recorded
August 1966
Genre
Pop, soul
Length
2:50
Label
Scepter
Songwriter(s)
Burt Bacharach and Hal David
Producer(s)
Burt Bacharach and Hal David
Dionne Warwick singles chronology
"Trains Boats and Planes"
(1966) "I Just Don't Know What to Do with Myself"
(1966) "The Beginning of Loneliness" / "Another Night"
(1966)
"Trains Boats and Planes"
(1966) "I Just Don't Know What to Do with Myself"
(1966) "The Beginning of Loneliness"/
"Another Night"
(1966)
Dionne Warwick recorded "I Just Don't Know What to Do with Myself" at Bell Sound Studios in August 1966 with Burt Bacharach producing; released that September the track is from her third album, Here Where There Is Love on Scepter Records. The song was a moderate success reaching #26 on the US Hot 100 and #20 on US Hot R&B with its strongest evident regional success in Detroit going Top Ten there.
After "Message to Michael" and "Trains and Boats and Planes" Warwick's "I Just Don't Know..." was her third consecutive single release comprising a previously recorded Bacharach/David song and Warwick's next Top 40 single "Alfie" would also fit in this category; three of the four appeared on Warwick's 1967 album Here Where There Is Love.
Formats and track listing[edit]
US 7" Vinyl Single
"I Just Don't Know What to Do With Myself" – 2:50
"In Between The Heartaches" – 2:50
Notes
"In Between The Heartaches" - Published By – Blue Seas Music, Inc. / JAC Music Co., Inc. (ASCAP)
"I Just Don't Know What To Do With Myself" - Published By – Quartet Music Inc. (BMI) Belinda (Canada Litd.)[4]
Charts[edit]
Chart (1966)
Peak
position
Canada Top Singles (RPM)[5] 36
US Cashbox Sales Chart[6] 24
US Billboard Hot 100[7] 26
US Billboard Hot R&B Singles[8] 20
Marcia Hines version[edit]
"I Just Don't Know What to Do with Myself"
Single by Marcia Hines
from the album Shining
A-side
"I Just Don't Know What to Do with Myself"
B-side
"Trilogy"
Released
1976
Format
single
Genre
Funk music, Soul music, Pop music
Length
3:08
Label
Wizard Records
Songwriter(s)
Burt Bacharach and Hal David
Producer(s)
Robie Porter
Marcia Hines singles chronology
"From the Inside"
(1975) "I Just Don't Know What to Do with Myself"
(1976) ""Don't Let the Grass Grow" / "You Gotta Let Go""
(1976)
"From the Inside"
(1975) "I Just Don't Know What to Do with Myself/ Trilogy"
(1976) ""Don't Let the Grass Grow" / "You Gotta Let Go""
(1976)
Marcia Hines recorded a version of the track and released it as the lead single from her second studio album, Shining (1976). The B-Side was "Trilogy" was included on her debut studio album, Marcia Shines (1975).
Track listing[edit]
7" Single (ZS-153)
Side A "I Just Don't Know What to Do with Myself" (Burt Bacharach/Hal David) - 3:08
Side B "Trilogy" (Robie Porter) - 4:20
Weekly charts[edit]
Chart (1975/76)
Position
Australian Kent Music Report[9]
10
Year-end charts[edit]
Chart (1976)
Position
Australian Kent Music Report[9]
23
See also[edit]
List of Top 25 singles for 1976 in Australia
White Stripes formats and track listing[edit]
"I Just Don't Know What to Do with Myself"
Single by The White Stripes
from the album Elephant
Released
September 2003
Format
CD, 7", DVD
Recorded
April 2003
Genre
Alternative rock, garage rock, blues rock
Length
2:46
Label
XL Recordings
Songwriter(s)
Burt Bacharach and Hal David
Producer(s)
Jack White
The White Stripes singles chronology
"Seven Nation Army"
(2003) "I Just Don't Know What to Do with Myself"
(2003) "The Hardest Button to Button"
(2003)
"Seven Nation Army"
(2003) "I Just Don't Know What to Do with Myself"
(2003) "The Hardest Button to Button"
(2003)
UK CD Promo
"I Just Don't Know What to Do with Myself" (BBC Radio 1 Evening Session) (Burt Bacharach, Hal David)
"Who's to Say..." (Dan Miller)
"Lafayette Blues" (BBC Radio 1 Evening Session)
"I'm Finding It Harder to Be a Gentleman "(Live on the John Peel Show)
US 7" Vinyl
"I Just Don't Know What to Do with Myself" (BBC Radio 1 Evening Session)
"Who's to Say..."
UK CD Single
"I Just Don't Know What to Do with Myself" (BBC Radio 1 Evening Session)
"Who's to Say"
"I'm Finding It Harder to Be a Gentleman" (Live on the John Peel Show)
UK DVD Single
"I Just Don't Know What to Do with Myself" (audio only)
"Lafayette Blues" (BBC Radio 1 Evening Session) (audio only)
"Black Math" (Masonic Temple Theatre, 16 April 2003)
Other versions[edit]
1964: A French version of the song by Michèle Vendôme and Claude Carrère, "Oui, il faut croire", was released by yéyé-singer Sheila.
1967: A band from Gadsden, Alabama The BLEUS, recorded a version of the song at Muscle Shoals Studio, with Eddie Hinton producing, released on the AMY label. The recording met with great success in the southeast U.S.
1968: An Italian translation by Franco Migliacci, "Se mi vuoi bene", was recorded by Patty Pravo. The original English version was also recorded by Cissy Houston in the same year.
1970: In October 1970 a Richard Perry-produced version of the song was released as the first solo single by Gary Puckett; primarily supported by Easy Listening radio, the track reached #61 on the Hot 100 in Billboard magazine whose Easy Listening chart afforded Puckett's "I Just Don't Know..." a #14 peak. Isaac Hayes released his version in the same year.
1972 the Dells.
1978 Elvis Costello & The Attractions; Demis Roussos. Costello later wrote of his version, "It was a measure of how backwards things were in 1977 that some people actually thought I was making a joke when The Attractions and I began performing "I Just Don't Know What to Do with Myself". I was not being ironic. I was being extremely literal."[10]
1980 The Photos (featuring Wendy Wu).
1991 the Stylistics.
1994 Linda Ronstadt.
1997 The soundtrack to My Best Friend's Wedding included a version by Nicky Holland. In the film, Cameron Diaz sings the song in a karaoke scene.
2003 Steve Tyrell (2003). In the same year, "I Just Don't Know What to Do with Myself" was released as a single by the American alternative rock band The White Stripes. It was the second single from their album Elephant. The video for the White Stripes version is a black-and-white video featuring Kate Moss who was pole dancing in her black bra and her black underwear, and was directed by Sofia Coppola.[11]
2006 The song was performed and included in the soundtrack for Shout! The Mod Musical.
2007 Trijntje Oosterhuis; Tina Arena.
2009 Scottish pop singer and songwriter Jimmy Somerville recorded the song for his album, Suddenly Last Summer.
2011 Ronan Keating recorded the song for his album, When Ronan Met Burt.
2012 Melanie C recorded the song for her musical-theatre-inspired album, Stages.
Although the recording year is unknown, a 2002 release of Smokey Robinson and the Miracles' Away We a Go-Go includes their version of "I Just Don't Know What to Do with Myself".
Bibliography[edit]
Serene Dominic. Burt Bacharach, song by song: the ultimate Burt Bacharach reference for fans. Schirmer Trade (New York NY) 2003. ISBN 0-8256-7280-5
References[edit]
Jump up ^ Dusty Springfield - UK Charts History. chartstats
Jump up ^ Ryan (bulion), Gary (15 June 2008). "Chart Positions Pre 1989 – Dusty Springfield". Australian Charts Portal. Hung Medien (Steffen Hung). Retrieved 3 July 2012.
Jump up ^ "dutchcharts.nl - Discografie Dusty Springfield". © 2006-2011 Hung Medien. Retrieved 2011-11-19.
Jump up ^ "I Just Don't Know"/"In Between The Heartaches" (7" Vinyl, Single) at Discogs
Jump up ^ RPM Archives. Dionne Warwick's Canada Singles History.
Jump up ^ Music: 1960's Songs Archives Files.
Jump up ^ Billboard Chart History. Dionne Warwick's Hot 100 History.
Jump up ^ Billboard Chart History. Dionne Warwick's R&B Singles History.
^ Jump up to: a b David Kent's "Australian Chart Book 1970-1992" Archived 5 March 2016 at Archive.is
Jump up ^ Elvis Costello (13 October 2015). Unfaithful Music & Disappearing Ink. Viking. p. 505. ISBN 978-0241003466.
Jump up ^ Lynn Hirschberg (31 August 2003). "The Coppola Smart Mob". NYTimes. Retrieved 16 February 2016.
External links[edit]
Lyrics of this song at MetroLyrics
White Stripes.net
[show]
v
t
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Dionne Warwick
Studio albums
Presenting Dionne Warwick (1963)
Anyone Who Had a Heart (1964)
Make Way for Dionne Warwick (1964)
The Sensitive Sound of Dionne Warwick (1965)
Here I Am (1965)
Here Where There Is Love (1966)
On Stage and in the Movies (1967)
The Windows of the World (1967)
The Magic of Believing (1968)
Dionne Warwick in Valley of the Dolls (1968)
Promises, Promises (1968)
Soulful (1969)
I'll Never Fall in Love Again (1970)
Very Dionne (1970)
Dionne (1972)
Just Being Myself (1973)
Then Came You (1975)
Track of the Cat (1975)
Love at First Sight (1977)
Dionne (1979)
No Night So Long (1980)
Friends in Love (1982)
Heartbreaker (1982)
How Many Times Can We Say Goodbye (1983)
Finder of Lost Loves (1985)
Friends (1985)
Reservations for Two (1987)
Dionne Sings Cole Porter (1990)
Friends Can Be Lovers (1993)
Aquarela do Brazil (1994)
Dionne Sings Dionne (1998)
Dionne Sings Dionne, Vol. 2 (2000)
My Favorite Time of the Year (2004)
My Friends & Me (2006)
Why We Sing (2008)
Only Trust Your Heart (2011)
Live albums
Dionne Warwick in Paris (1966)
Hot! Live and Otherwise (1981)
Christmas in Vienna II (1993)
Compilation albums
The Love Collection (2008)
Singles
"Don't Make Me Over"
"Wishin' and Hopin'"
"Anyone Who Had a Heart"
"Walk On By"
"You'll Never Get to Heaven (If You Break My Heart)"
"A House Is Not a Home"
"Reach Out for Me"
"Who Can I Turn To?"
"Are You There (With Another Girl)"
"Message to Michael"
"I Just Don't Know What to Do with Myself"
"Alfie"
"The Windows of the World"
"I Say a Little Prayer"
"(Theme from) Valley of the Dolls"
"Do You Know the Way to San Jose"
"(There's) Always Something There to Remind Me"
"This Girl's in Love with You"
"You've Lost That Lovin' Feeling"
"I'll Never Fall in Love Again"
"Make It Easy on Yourself"
"The Good Life"
"Then Came You" (with Spinners)
"By the Time I Get to Phoenix/I Say a Little Prayer" (with Isaac Hayes)
"Only Love Can Break a Heart"
"I'll Never Love This Way Again"
"Déjà Vu"
"After You"
"No Night So Long"
"Easy Love"
"Heartbreaker"
"All the Love in the World"
"Run to Me" (with Barry Manilow)
"That's What Friends Are For" (with Elton John, Gladys Knight & Stevie Wonder)
"Love Power" (with Jeffrey Osborne)
"It's All Over" (with Dieter Bohlen as Blue System)
"What the World Needs Now Is Love" (with TheHipHopNationUnited)
Record labels
Arista
Concord
Rhino
River North
Scepter
Warner Bros.
Related articles
Discography
Solid Gold (TV series)
Book:Dionne Warwick
[show]
v
t
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Marcia Hines
Discography
Studio albums
Marcia Shines (1975)
Shining (1976)
Ladies and Gentlemen (1977)
Ooh Child (1979)
Take It from the Boys (1981)
Jokers and Queens (with Jon English) (1982)
Love Sides (1983)
Right Here and Now (1994)
Time of Our Lives (1999)
Hinesight (2004)
Discothèque (2006)
Life (2007)
Marcia Sings Tapestry (2010)
Amazing (2014)
Live albums
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Stoma
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Not to be confused with Stroma.
For natural and surgically created body openings, see Stoma (medicine).
Stoma in a tomato leaf shown via colorized scanning electron microscope image
A stoma in cross section
The underside of a leaf. In this species (Tradescantia zebrina) the guard cells of the stomata are green because they contain chlorophyll while the epidermal cells are chlorophyll-free and contain red pigments.
In botany, a stoma (plural "stomata"), also called a stomate (plural "stomates")[1] (from Greek στόμα, "mouth"[2]), is a pore, found in the epidermis of leaves, stems, and other organs, that facilitates gas exchange. The pore is bordered by a pair of specialized parenchyma cells known as guard cells that are responsible for regulating the size of the stomatal opening.
The term is usually used collectively to refer to the entire stomatal complex, consisting of the paired guard cells and the pore itself, which is referred to as the stomatal aperture.[3] Air enters the plant through these openings by gaseous diffusion, and contains carbon dioxide and oxygen, which are used in photosynthesis and respiration, respectively. Oxygen produced as a by-product of photosynthesis diffuses out to the atmosphere through these same openings. Also, water vapor diffuses through the stomata into the atmosphere in a process called transpiration.
Stomata are present in the sporophyte generation of all land plant groups except liverworts. In vascular plants the number, size and distribution of stomata varies widely. Dicotyledons usually have more stomata on the lower surface of the leaves than the upper surface. Monocotyledons such as onion, oat and maize may have about the same number of stomata on both leaf surfaces.[4]:5 In plants with floating leaves, stomata may be found only on the upper epidermis and submerged leaves may lack stomata entirely. Most tree species have stomata only on the lower leaf surface.[5] Leaves with stomata on both the upper and lower leaf are called amphistomatous leaves; leaves with stomata only on the lower surface are hypostomatous, and leaves with stomata only on the upper surface are epistomatous or hyperstomatous.[5] Size varies across species, with end-to-end lengths ranging from 10 to 80 µm and width ranging from a few to 50 µm.[6]
Contents
[hide]
1 Function
1.1 CO2 gain and water loss
1.2 Alternative approaches
1.3 CAM plants
1.4 Opening and closure
1.5 Inferring stomatal behavior from gas exchange
2 Evolution
3 Development
4 Stomatal crypts
5 Stomata as pathogenic pathways
6 Stomata and climate change
6.1 Response of stomata to environmental factors
6.2 Future adaptations during climate change
6.3 Agricultural implications
7 References
8 External links
Function[edit]
Electron micrograph of a stoma from a Brassica chinensis (Bok Choy) leaf
CO2 gain and water loss[edit]
Carbon dioxide, a key reactant in photosynthesis, is present in the atmosphere at a concentration of about 400 ppm. Most plants require the stomata to be open during daytime. The air spaces in the leaf are saturated with water vapour, which exits the leaf through the stomata; this is known as transpiration. Therefore, plants cannot gain carbon dioxide without simultaneously losing water vapour.[7]
Alternative approaches[edit]
Ordinarily, carbon dioxide is fixed to ribulose-1,5-bisphosphate (RuBP) by the enzyme RuBisCO in mesophyll cells exposed directly to the air spaces inside the leaf. This exacerbates the transpiration problem for two reasons: first, RuBisCo has a relatively low affinity for carbon dioxide, and second, it fixes oxygen to RuBP, wasting energy and carbon in a process called photorespiration. For both of these reasons, RuBisCo needs high carbon dioxide concentrations, which means wide stomatal apertures and, as a consequence, high water loss.
Narrower stomatal apertures can be used in conjunction with an intermediary molecule with a high carbon dioxide affinity, PEPcase (Phosphoenolpyruvate carboxylase). Retrieving the products of carbon fixation from PEPCase is an energy-intensive process, however. As a result, the PEPCase alternative is preferable only where water is limiting but light is plentiful, or where high temperatures increase the solubility of oxygen relative to that of carbon dioxide, magnifying RuBisCo's oxygenation problem.
CAM plants[edit]
C3 and C4 plants(1) stomata stay open all day and close at night. CAM plants(2) stomata open during the morning and close slightly at noon and then open again in the morning.
A group of mostly desert plants called "CAM" plants (Crassulacean acid metabolism, after the family Crassulaceae, which includes the species in which the CAM process was first discovered) open their stomata at night (when water evaporates more slowly from leaves for a given degree of stomatal opening), use PEPcarboxylase to fix carbon dioxide and store the products in large vacuoles. The following day, they close their stomata and release the carbon dioxide fixed the previous night into the presence of RuBisCO. This saturates RuBisCO with carbon dioxide, allowing minimal photorespiration. This approach, however, is severely limited by the capacity to store fixed carbon in the vacuoles, so it is preferable only when water is severely limiting.
Opening and closure[edit]
For more details on this topic, see Guard cell.
Opening and closing of stoma.
However, most plants do not have the aforementioned facility and must therefore open and close their stomata during the daytime, in response to changing conditions, such as light intensity, humidity, and carbon dioxide concentration. It is not entirely certain how these responses work. However, the basic mechanism involves regulation of osmotic pressure.
When conditions are conducive to stomatal opening (e.g., high light intensity and high humidity), a proton pump drives protons (H+) from the guard cells. This means that the cells' electrical potential becomes increasingly negative. The negative potential opens potassium voltage-gated channels and so an uptake of potassium ions (K+) occurs. To maintain this internal negative voltage so that entry of potassium ions does not stop, negative ions balance the influx of potassium. In some cases, chloride ions enter, while in other plants the organic ion malate is produced in guard cells. This increase in solute concentration lowers the water potential inside the cell, which results in the diffusion of water into the cell through osmosis. This increases the cell's volume and turgor pressure. Then, because of rings of cellulose microfibrils that prevent the width of the guard cells from swelling, and thus only allow the extra turgor pressure to elongate the guard cells, whose ends are held firmly in place by surrounding epidermal cells, the two guard cells lengthen by bowing apart from one another, creating an open pore through which gas can move.[8]
When the roots begin to sense a water shortage in the soil, abscisic acid (ABA) is released.[9] ABA binds to receptor proteins in the guard cells' plasma membrane and cytosol, which first raises the pH of the cytosol of the cells and cause the concentration of free Ca2+ to increase in the cytosol due to influx from outside the cell and release of Ca2+ from internal stores such as the endoplasmic reticulum and vacuoles.[10] This causes the chloride (Cl−) and organic ions to exit the cells. Second, this stops the uptake of any further K+ into the cells and, subsequently, the loss of K+. The loss of these solutes causes an increase in water potential, which results in the diffusion of water back out of the cell by osmosis. This makes the cell plasmolysed, which results in the closing of the stomatal pores.
Guard cells have more chloroplasts than the other epidermal cells from which guard cells are derived. Their function is controversial.[11][12]
Inferring stomatal behavior from gas exchange[edit]
The degree of stomatal resistance can be determined by measuring leaf gas exchange of a leaf. The transpiration rate is dependent on the diffusion resistance provided by the stomatal pores, and also on the humidity gradient between the leaf's internal air spaces and the outside air. Stomatal resistance (or its inverse, stomatal conductance) can therefore be calculated from the transpiration rate and humidity gradient. This allows scientists to investigate how stomata respond to changes in environmental conditions, such as light intensity and concentrations of gases such as water vapor, carbon dioxide, and ozone.[13] Evaporation (E) can be calculated as;[14]
E = ( e i − e a ) / P r {\displaystyle E=(e_{i}-e_{a})/Pr}
where ei and ea are the partial pressures of water in the leaf and in the ambient air, respectively, P is atmospheric pressure, and r is stomatal resistance. The inverse of r is conductance to water vapor (g), so the equation can be rearranged to;[14]
E = ( e i − e a ) g / P {\displaystyle E=(e_{i}-e_{a})g/P}
and solved for g;[14]
g = E P / ( e i − e a ) {\displaystyle g=EP/(e_{i}-e_{a})}
Photosynthetic CO2 assimilation (A) can be calculated from
A = ( C a − C i ) g / 1.6 P {\displaystyle A=(C_{a}-C_{i})g/1.6P}
where Ca and Ci are the atmospheric and sub-stomatal partial pressures of CO2, respectively. The rate of evaporation from a leaf can be determined using a photosynthesis system. These scientific instruments measure the amount of water vapour leaving the leaf and the vapor pressure of the ambient air. Photosynthetic systems may calculate water use efficiency (A/E), g, intrinsic water use efficiency (A/g), and Ci. These scientific instruments are commonly used by plant physiologists to measure CO2 uptake and thus measure photosynthetic rate.[15] [16]
Evolution[edit]
Tomato stoma observed through immersion oil
There is little evidence of the evolution of stomata in the fossil record, but they had appeared in land plants by the middle of the Silurian period.[17] They may have evolved by the modification of conceptacles from plants' alga-like ancestors.[18] However, the evolution of stomata must have happened at the same time as the waxy cuticle was evolving – these two traits together constituted a major advantage for early terrestrial plants.[citation needed]
Development[edit]
There are three major epidermal cell types which all ultimately derive from the outermost (L1) tissue layer of the shoot apical meristem, called protodermal cells: trichomes, pavement cells and guard cells, all of which are arranged in a non-random fashion.
An asymmetrical cell division occurs in protodermal cells resulting in one large cell that is fated to become a pavement cell and a smaller cell called a meristemoid that will eventually differentiate into the guard cells that surround a stoma. This meristemoid then divides asymmetrically one to three times before differentiating into a guard mother cell. The guard mother cell then makes one symmetrical division, which forms a pair of guard cells.[19] Cell division is inhibited in some cells so there is always at least one cell between stomata.[20]
Stomatal patterning is controlled by the interaction of many signal transduction components such as EPF (Epidermal Patterning Factor), ERL (ERecta Like) and YODA (a putative MAP kinase kinase kinase).[20] Mutations in any one of the genes which encode these factors may alter the development of stomata in the epidermis.[20] For example, a mutation in one gene causes more stomata that are clustered together, hence is called Too Many Mouths (TMM).[19] Whereas, disruption of the SPCH (SPeecCHless) gene prevents stomatal development all together.[20] Activation of stomatal production can occur by the activation of EPF1, which activates TMM/ERL, which together activate YODA. YODA inhibits SPCH, causing SPCH activity to decrease, allowing for asymmetrical cell division that initiates stomata formation.[20][21] Stomatal development is also coordinated by the cellular peptide signal called stomagen, which signals the inhibition of the SPCH, resulting in increased number of stomata.[22]
Environmental and hormonal factors can affect stomatal development. Light increases stomatal development in plants; while, plants grown in the dark have a lower amount of stomata. Auxin represses stomatal development by affecting their development at the receptor level like the ERL and TMM receptors. However, a low concentration of auxin allows for equal division of a guard mother cell and increases the chance of producing guard cells.[23]
Stomatal crypts[edit]
Stomatal crypts are sunken areas of the leaf epidermis which form a chamber-like structure that contains one or more stomata and sometimes trichomes or accumulations of wax. Stomatal crypts can be an adaption to drought and dry climate conditions when the stomatal crypts are very pronounced. However, dry climates are not the only places where they can be found. The following plants are examples of species with stomatal crypts or antechambers: Nerium oleander, conifers, and Drimys winteri which is a species of plant found in the cloud forest.[24]
Stomata as pathogenic pathways[edit]
Stomata are obvious holes in the leaf by which, as was presumed for a while, pathogens can enter unchallenged. However, it has been recently shown that stomata do in fact sense the presence of some, if not all, pathogens. However, with the virulent bacteria applied to Arabidopsis plant leaves in the experiment, the bacteria released the chemical coronatine, which forced the stomata open again within a few hours.[25]
Stomata and climate change[edit]
Response of stomata to environmental factors[edit]
Photosynthesis, plant water transport (xylem) and gas exchange are regulated by stomatal function which is important in the functioning of plants.[26] Stomatal density and aperture (length of stomata) varies under a number of environmental factors such as atmospheric CO2 concentration, light intensity, air temperature and photoperiod (daytime duration). [27][28]
Decreasing stomatal density is one way plants have responded to the increase in concentration of atmospheric CO2 ([CO2]atm).[29] Although changes in [CO2]atm response is the least understood mechanistically, this stomatal response has begun to plateau where it is soon expected to impact transpiration and photosynthesis processes in plants.[26][30]
Future adaptations during climate change[edit]
It is expected for [CO2]atm to reach 500–1000 ppm by 2100.[26] 96% of the past 400 000 years experienced below 280 ppm CO2 levels. From this figure, it is highly probable that genotypes of today’s plants diverged from their pre-industrial relative. [26]
The gene HIC (high carbon dioxide) encodes a negative regulator for the development of stomata in plants.[31] Research into the HIC gene using Arabidopsis thaliana found no increase of stomatal development in the dominant allele, but in the ‘wild type’ recessive allele showed a large increase, both in response to rising CO2 levels in the atmosphere.[31] These studies imply the plants response to changing CO2 levels is largely controlled by genetics.
Agricultural implications[edit]
In the face of ecological contingencies such as increasing temperatures, changes in rainfall patterns, long term climate change, and biotic influences of human management interventions, it is expected to reduce the production and quality of food and have a negative impact on agricultural production.[28][29]
The CO2 fertiliser effect has been greatly overestimated during Free-Air Carbon dioxide Enrichment (FACE) experiments where results show increased CO2 levels in the atmosphere enhances photosynthesis, reduce transpiration, and increase water use efficiency (WUE).[29] Increased biomass is one of the effects with simulations from experiments predicting a 5–20% increase in crop yields at 550 ppm of CO2.[32] Rates of leaf photosynthesis were shown to increase by 30–50% in C3 plants, and 10–25% in C4 under doubled CO2 levels.[32] The existence of a feedback mechanism results a phenotypic plasticity in response to [CO2]atm that may have been an adaptive trait in the evolution of plant respiration and function.[26][28]
Predicting how stomata perform during adaptation is useful for understanding the productivity of plant systems for both natural and agricultural systems.[27] Plant breeders and farmers are beginning to work together using evolutionary and participatory plant breeding to find the best suited species such as heat and drought resistant crop varieties that could naturally evolve to the change in the face of food security challenges.[29]
References[edit]
Jump up ^ "Living Environment—Regents High school examination" (PDF). January 2011 Regents. NYSED. Retrieved 15 June 2013.
Jump up ^ στόμα, Henry George Liddell, Robert Scott, A Greek-English Lexicon, on Perseus
Jump up ^ Esau, K. (1977). Anatomy of Seed Plants. Wiley and Sons. p. 88. ISBN 0-471-24520-8.
Jump up ^ Weyers, J. D. B.; Meidner, H. (1990). Methods in stomatal research. Longman Group UK Ltd. ISBN 0582034833.
^ Jump up to: a b Willmer, Colin; Fricker, Mark (1996). Stomata. Springer. p. 16. doi:10.1007/978-94-011-0579-8.
Jump up ^ Fricker, M.; Willmer, C. (2012). Stomata. Springer Netherlands. p. 18. ISBN 978-94-011-0579-8. Retrieved 15 June 2016.
Jump up ^ Debbie Swarthout and C.Michael Hogan. 2010. Stomata. Encyclopedia of Earth. National Council for Science and the Environment, Washington DC
Jump up ^ N. S. CHRISTODOULAKIS; J. MENTI; B. GALATIS (January 2002). "Structure and Development of Stomata on the Primary Root of Ceratonia siliqua L". Annals of Botany. 89 (1): 23–29. doi:10.1093/aob/mcf002. PMID 12096815.
Jump up ^ C. L. Trejo; W. J. Davies; LdMP. Ruiz (1993). "Sensitivity of Stomata to Abscisic Acid (An Effect of the Mesophyll)". Plant Physiology. 102 (2): 497–502. doi:10.1104/pp.102.2.497. PMC 158804  . PMID 12231838.
Jump up ^ Petra Dietrich; Dale Sanders; Rainer Hedrich (October 2001). "The role of ion channels in light-dependent stomatal opening". Journal of Experimental Botany. 52 (363): 1959–1967. doi:10.1093/jexbot/52.363.1959. PMID 11559731.
Jump up ^ "Guard Cell Photosynthesis". Retrieved 2015-10-04.
Jump up ^ Eduardo Zeiger; Lawrence D. Talbott; Silvia Frechilla; Alaka Srivastava; Jianxin Zhu (March 2002). "The Guard Cell Chloroplast: A Perspective for the Twenty-First Century". New Phytologist. 153 (3 Special Issue: Stomata): 415–424. doi:10.1046/j.0028-646X.2001.NPH328.doc.x.
Jump up ^ Hopkin, Michael (2007-07-26). "Carbon sinks threatened by increasing ozone". Nature. 448 (7152): 396–397. Bibcode:2007Natur.448..396H. doi:10.1038/448396b. PMID 17653153.
^ Jump up to: a b c "Calculating Important Parameters in Leaf Gas Exchange". Plant Physiology Online. Sinauer. Retrieved 2013-02-24.
Jump up ^ Waichi Agata; Yoshinobu Kawamitsu; Susumu Hakoyama; Yasuo Shima (January 1986). "A system for measuring leaf gas exchange based on regulating vapour pressure difference". Photosynthesis Research. 9 (3): 345–357. doi:10.1007/BF00029799. ISSN 1573-5079. PMID 24442366. Retrieved May 6, 2010.
Jump up ^ Portable Gas Exchange Fluorescence System GFS-3000. Handbook of Operation (PDF), March 20, 2013
Jump up ^ D. Edwards, H. Kerp; Hass, H. (1998). "Stomata in early land plants: an anatomical and ecophysiological approach". Journal of Experimental Botany. 49 (Special Issue): 255–278. doi:10.1093/jxb/49.Special_Issue.255.
Jump up ^ Krassilov, Valentin A. (2004). "Macroevolutionary events and the origin of higher taxa". In Wasser, Solomon P. Evolutionary theory and processes : modern horizons : papers in honour of Eviatar Nevo. Dordrecht: Kluwer Acad. Publ. pp. 265–289. ISBN 1-4020-1693-X.
^ Jump up to: a b Bergmann, Dominique C.; Lukowitz, Wolfgang; Somerville, Chris R.; Lukowitz, W; Somerville, CR (4 July 2004). "Stomatal Development and Pattern Controlled by a MAPKK Kinase". Science. 304 (5676): 1494–1497. doi:10.1126/science.1096014. PMID 15178800.
^ Jump up to: a b c d e Pillitteri, Lynn Jo; Dong, Juan (2013-06-06). "Stomatal Development in Arabidopsis". The Arabidopsis Book / American Society of Plant Biologists. 11: e0162. doi:10.1199/tab.0162. ISSN 1543-8120. PMC 3711358  . PMID 23864836.
Jump up ^ Casson, Stuart A; Hetherington, Alistair M (2010-02-01). "Environmental regulation of stomatal development". Current Opinion in Plant Biology. 13 (1): 90–95. doi:10.1016/j.pbi.2009.08.005.
Jump up ^ Sugano, Shigeo S.; Shimada, Tomoo; Imai, Yu; Okawa, Katsuya; Tamai, Atsushi; Mori, Masashi; Hara-Nishimura, Ikuko (2010-01-14). "Stomagen positively regulates stomatal density in Arabidopsis". Nature. 463 (7278): 241–244. doi:10.1038/nature08682. ISSN 0028-0836.
Jump up ^ Balcerowicz, M.; Ranjan, A.; Rupprecht, L.; Fiene, G.; Hoecker, U. (2014). "Auxin represses stomatal development in dark-grown seedling via Aux/IAA proteins". Development. 141: 3165–3176.
Jump up ^ Roth-Nebelsick, A.; Hassiotou, F.; Veneklaas, E. J (2009). "Stomatal crypts have small effects on transpiration: A numerical model analysis". Plant Physiology. 151: 2018–2027. doi:10.1104/pp.109.146969.
Jump up ^ Maeli Melotto; William Underwood; Jessica Koczan; Kinya Nomura; Sheng Yang He (September 2006). "Plant Stomata Function in Innate Immunity against Bacterial Invasion". Cell. 126 (5): 969–980. doi:10.1016/j.cell.2006.06.054. PMID 16959575.
^ Jump up to: a b c d e Rico, C; Pittermann, J; Polley, HW; Aspinwall, MJ; Fay, PA (2013). "The effect of subambient to elevated atmospheric CO2 concentration on vascular function in Helianthus annuus: implications for plant response to climate change". New Phytologist. 199: 956–965. doi:10.1111/nph.12339.
^ Jump up to: a b Buckley, TN; Mott, KA (2013). "Modelling stomatal conductance in response to environmental factors". Plant, Cell and Environment. 36: 1691–1699. doi:10.1111/pce.12140.
^ Jump up to: a b c Rogiers, SY; Hardie, WJ; Smith, JP (2011). "Stomatal density of grapevine leaves (Vitis Vinifera L.) responds to soil temperature and atmospheric carbon dioxide". Australian Journal of Grape and Wine Research. 17: 147–152. doi:10.1111/j.1755-0238.2011.00124.x.
^ Jump up to: a b c d Ceccarelli, S; Grando, S; Maatougui, M; Michael, M; Slash, M; Haghparast, R; Rahmanian, M; Taheri, A; Al-Yassin, A; Benbelkacem, A; Labdi, M; Mimoun, H; Nachit, M (2010). "'Plant breeding and climate changes'". The Journal of Agricultural Science. 148 (6): 627–637. doi:10.1017/s0021859610000651.
Jump up ^ Serna, L; Fenoll, C (2000). "Coping with human CO2 emissions". Nature. 408: 656–657. doi:10.1038/35047202.
^ Jump up to: a b Gray, J; Holroyd, G; van der Lee, F; Bahrami, A; Sijmons, P; Woodward, F; Schuch, W; Hetherington, A (2000). "The HIC signalling pathway links CO2 perception to stomatal development". Nature. 408: 713–716.
^ Jump up to: a b Tubiello, FN; Soussana, J-F; Howden, SM (2007). "Crop and pasture response to climate change". Proceedings of the National Academy of Sciences of the United States of America. 104 (50): 19686–19690. doi:10.1073/pnas.0701728104.
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