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2018 Winter Olympics
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"PyeongChang 2018" redirects here. For the Winter Paralympics, see 2018 Winter Paralympics.
XXIII Olympic Winter Games
Host city
Pyeongchang, South Korea
Motto
Passion. Connected.
Korean: 하나된 열정. (Hanadoen Yeoljeong)
Nations participating
90 (estimated)
Events
102 in 7 sports (15 disciplines)
Opening ceremony
9 February (4 months from now)
Closing ceremony
25 February
Stadium
Pyeongchang Olympic Stadium
Winter
< Sochi 2014
Beijing 2022 >
Summer
< Rio de Janeiro 2016
Tokyo 2020 >
Pyeongchang Winter Olympics
Hangul
평창 동계 올림픽
Hanja
平昌 冬季 올림픽
Revised Romanization
Pyeongchang Donggye Ollimpik
McCune–Reischauer
P'yŏngch'ang Donggye Ollimp'ik
This article contains Korean text. Without proper rendering support, you may see question marks, boxes, or other symbols instead of Hangul and Hanja.
Part of a series on
2018 Winter Olympics[show]
Bid process (bid details)
Venues
Torch relay
Mascots
Broadcasters
Opening ceremony (flag bearers)
Chronological summary
Medal table (medalists)
World and Olympic records
Concerns and controversies
Closing ceremony (flag bearers)
Paralympics
IOC
KOC
POCOG
v
t
e
The 2018 Winter Olympics, officially known as the XXIII Olympic Winter Games (French: Les XXIIIeme Jeux olympiques d'hiver; Hangul: 평창 동계 올림픽; RR: Pyeongchang Donggye Ollimpik), and commonly known as PyeongChang 2018,[1] [pʰjʌŋ.tɕʰaŋ] is a major international multi-sport event scheduled to take place from 9 to 25 February 2018 in Pyeongchang County, South Korea.
The elected host city was announced on 6 July 2011 by the International Olympic Committee (IOC) after the 123rd IOC Session in Durban, South Africa. Pyeongchang won its bid on the first round of voting, receiving more votes than both Munich, Germany and Annecy, France.
The 2018 Olympics will be the second Olympic Games held in South Korea, after the 1988 Summer Olympics held in Seoul, and will be South Korea's first Winter Games. Pyeongchang will be the third Asian city to host the Winter Games after Sapporo, Japan (1972), and Nagano, Japan (1998).[2]
Contents
[hide]
1 Bidding
1.1 Host City Election
2 Tickets
3 Preparations
4 Venues
4.1 Pyeongchang Mountain cluster
4.1.1 Alpensia Sports Park
4.1.2 Stand-alone venues
4.2 Gangneung Coastal cluster
5 Sports
6 Participating National Olympic Committees
7 Calendar
8 Marketing
8.1 Branding
8.2 Mascots
8.3 Sponsors
9 Broadcasting rights
10 Concerns and controversies
10.1 Russian Doping
10.2 Security
11 See also
12 Notes and references
13 External links
Bidding[edit]
Main article: Bids for the 2018 Winter Olympics
Pyeongchang bid to host both the 2010 and 2014 Winter Olympic Games but lost in the final rounds of voting by three and four votes respectively. Pyeongchang won its bid for the 2018 Winter Olympic Games in the first round of voting, receiving 63 of the 95 votes cast, giving it the majority required to be elected host city.
Munich also launched a bid to host these Games. Prior to Beijing's successful 2022 Winter Olympics bid, Munich would have become the first city to host both the Winter and the Summer Games, having previously hosted the 1972 Summer Olympics, but received 25 votes. Annecy launched a bid, but failed to secure public support from local citizens. Their bid received seven votes.
Host City Election[edit]
PyeongChang was elected as the host city at the 123rd IOC Session in Durban. Forty-eight votes were needed for selection and PyeongChang was elected after just one round of voting.
2018 Winter Olympics bidding results
City
Nation
Votes
Pyeongchang South Korea 63
Munich Germany 25
Annecy France 7
Tickets[edit]
The ticket prices for the 2018 Winter Olympics were announced in April 2016 and went on sale in October 2016, ranging from ₩20,000 approx. $18 to ₩900,000 ($803). Tickets for the opening and closing ceremonies will range from ₩220,000 ($196) to ₩1.5 million ($1,338). The exact prices were determined through market research; around 50% of the tickets are due to cost about ₩80,000 ($71) or less, and tickets in sports that are relatively unknown in the region, such as biathlon and luge, will be made cheaper in order to encourage attendance. By contrast, figure skating and the Men's hockey gold medal game carry the most expensive tickets of the Games.[3]
Preparations[edit]
On 5 August 2011, the International Olympic Committee (IOC) announced the formation of the Pyeongchang 2018 Coordination Commission.[4][5] On 4 October 2011, it was announced that the Organizing Committee for the 2018 Winter Olympics will be headed by Kim Jin-sun. The Pyeongchang Organizing Committee for the 2018 Olympic & Paralympic Winter Games (POCOG) was launched at its inaugural assembly on 19 October 2011. The first tasks of the organizing committee was to put together a master plan for the games as well as forming a design for the venues.[6] The IOC Coordination Commission for the 2018 Winter Olympics made their first visit to Pyeongchang in March 2012. By then, construction was already underway on the Olympic Village.[7][8] In June 2012, construction began on a high-speed rail line that will connect Pyeongchang to Seoul.[9]
Olympic venues 2018
The International Paralympic Committee met with the Pyeongchang 2018 organizing committee for an orientation in July 2012.[10] Then-IOC President Jacques Rogge visited Pyeongchang for the first time in February 2013.[11]
On 27 June 2014 the PyeongChang Olympic Committee announced their mascot selection contest.[12] The contest ran from 15 September 2014 to 30 September 2014. The 2013 Special Olympics World Winter Games were held in Pyeongchang.
The Pyeongchang Organizing Committee for the 2018 Olympic & Paralympic Winter Games created Pyeongchang WINNERS in 2014 by recruiting university students living in South Korea to spread awareness of the Olympic Games through social networking services and news articles.[13]
Venues[edit]
Main article: Venues of the 2018 Winter Olympics and Paralympics
Dragon Valley Ski Resort
Pyeongchang Mountain cluster[edit]
Alpensia Sports Park[edit]
The Alpensia Resort in Daegwallyeong-myeon will be the focus of the 2018 Pyeongchang Winter Olympics.[14][15]
Pyeongchang Olympic Stadium – opening and closing ceremonies[16]
Alpensia Ski Jumping Centre – ski jumping, Nordic combined, snowboarding (big air)
Alpensia Biathlon Centre – biathlon
Alpensia Cross-Country Centre – cross-country skiing, Nordic combined
Alpensia Sliding Centre – luge, bobsleigh, and skeleton
Olympic Village
Yongpyong Alpine Centre – alpine skiing (slalom, giant slalom)
Stand-alone venues[edit]
Bokwang Snow Park – freestyle skiing and snowboard
Jeongseon Alpine Centre – alpine skiing (downhill, super-G, and combined)
Gangneung Coastal cluster[edit]
The coastal cluster is located in the city of Gangneung. The Gangneung Olympic Park will include the following four venues:
Gangneung Hockey Centre – ice hockey (men competition)
Gangneung Curling Centre – curling
Gangneung Oval[16] – speed skating
Gangneung Ice Arena – short track speed skating and figure skating
In addition, a stand-alone venue is located on the grounds of Catholic Kwandong University:
Kwandong Hockey Centre – ice hockey (women competition)
Sports[edit]
The 2018 Winter Olympics will feature 102 events in 15 sports. Four new disciplines in existing sports will be introduced in Pyeongchang, including big air snowboarding (which will replace the parallel slalom), mixed doubles curling, mass start speed skating, and mixed team alpine skiing.[17]
For the first time since 1998, the National Hockey League will not provide accommodations (including a break in the season for all teams during the Olympics) to allow its players to participate in the Men's ice hockey tournament. The NHL's decision stemmed from demands for the IOC to cover the cost of insuring the NHL players that participate in the Games. Although it did pay to insure NHL players in Sochi, the IOC was unwilling to do so for Pyeongchang, and was concerned that the NHL's demand could set a precedent for other professional sports bodies to follow in the future. NHL Commissioner Gary Bettman added that the IOC didn't allow the NHL to promote the involvement of its players in the Olympics.[18][19][20] The NHL has yet to announce if it will implement any specific measures to penalize players who participate in the Olympics.[21]
Numbers in parentheses indicate the number of medal events contested in each sports discipline.
Alpine skiing (11) (details)
Biathlon (11) (details)
Bobsleigh (3) (details)
Cross-country skiing (12) (details)
Curling (3) (details)
Figure skating (5) (details)
Freestyle skiing (10) (details)
Ice hockey (2) (details)
Luge (4) (details)
Nordic combined (3) (details)
Short track speed skating (8) (details)
Skeleton (2) (details)
Ski jumping (4) (details)
Snowboarding (10) (details)
Speed skating (14) (details)
Participating National Olympic Committees[edit]
A total of 83 nations have qualified at least one athlete so far.
Eritrea, Kosovo and Malaysia are scheduled to make their Winter Olympics debut if they decide to participate.
If North Korea qualifies for the Games, they will be allowed to cross the DMZ into South Korea.[22]
Russia's status remains unclear due to the state-sponsored doping program scandal.[23]
[hide]Participating National Olympic Committees[24][25][26][27]
Albania
Andorra
Argentina
Armenia
Australia
Austria
Azerbaijan
Belarus
Belgium
Bermuda
Bosnia and Herzegovina
Bolivia
Brazil
Bulgaria
Canada
Cayman Islands
Chile
China
Colombia
Croatia
Cyprus
Czech Republic
Denmark
Dominica
Eritrea
Estonia
Finland
France
Georgia
Germany
Great Britain
Greece
Hungary
Iceland
Iran
Ireland
Israel
Italy
Japan
Kazakhstan
Kenya
Kosovo
Kyrgyzstan
Latvia
Lebanon
Liechtenstein
Lithuania
Luxembourg
Macedonia
Madagascar
Malaysia
Malta
Mexico
Monaco
Mongolia
Montenegro
Morocco
Netherlands
New Zealand
Norway
Pakistan
Peru
Poland
Portugal
Puerto Rico
Romania
Russia
San Marino
Serbia
Slovakia
Slovenia
South Africa
South Korea (host)
Spain
Sweden
Switzerland
Thailand
Timor-Leste
Togo
Turkey
Ukraine
United States
Uzbekistan
Calendar[edit]
All dates are KST (UTC+9)

OC Opening ceremony ● Event competitions 1 Event finals EG Exhibition gala CC Closing ceremony
February
8th
Thu
9th
Fri
10th
Sat
11th
Sun
12th
Mon
13th
Tue
14th
Wed
15th
Thu
16th
Fri
17th
Sat
18th
Sun
19th
Mon
20th
Tue
21st
Wed
22nd
Thu
23rd
Fri
24th
Sat
25th
Sun
Events
Ceremonies OC CC N/A
Alpine skiing 1 1 1 1 1 1 1 1 1 1 1 11
Biathlon 1 1 2 1 1 1 1 1 1 1 11
Bobsleigh ● 1 ● 1 ● 1 3
Cross country skiing 1 1 2 1 1 1 1 2 1 1 12
Curling ● ● ● ● ● 1 ● ● ● ● ● ● ● ● ● ● 1 1 3
Figure skating ● ● 1 ● 1 ● 1 ● 1 ● 1 EG 5
Freestyle skiing ● 1 1 ● 1 1 2 ● 1 1 1 1 10
Ice hockey ● ● ● ● ● ● ● ● ● ● ● ● 1 ● ● 1 2
Luge ● 1 ● 1 1 1 4
Nordic combined 1 1 1 3
Short track speed skating 1 1 2 1 3 8
Skeleton ● 1 1 2
Ski jumping ● 1 1 ● 1 1 4
Snowboarding ● 1 1 1 1 1 1 ● ● ● 1 3 10
Speed skating 1 1 1 1 1 1 1 1 1 2 1 2 14
Total events
5
7
8
8
6
7
5
9
6
3
5
7
8
6
8
4
102
Cumulative total
5
12
20
28
34
41
46
55
61
64
69
76
84
90
98
102
February
8th
Thu
9th
Fri
10th
Sat
11th
Sun
12th
Mon
13th
Tue
14th
Wed
15th
Thu
16th
Fri
17th
Sat
18th
Sun
19th
Mon
20th
Tue
21st
Wed
22nd
Thu
23rd
Fri
24th
Sat
25th
Sun
102 events

Marketing[edit]
Gold medal of the 2018 Olympics
Branding[edit]
The emblem for the Games was unveiled on 3 May 2013. It is a stylized representation of the hangul letters ㅍ p and ㅊ ch, being the initial sounds of 평창 Pyeongchang. Additionally the left symbol is said to represent the Korean philosophical triad of heaven, earth and humanity (Korean: 천지인 cheon-ji-in), and the right symbol to represent a crystal of ice.[28]
The name of the host city, Pyeongchang, has been intentionally styled in CamelCase as "PyeongChang" on the emblem and in official materials, rather than "Pyeongchang". This has been done to alleviate potential confusion with Pyongyang, the similarly-named capital of neighbouring North Korea. Gangwon Governor Choi Moon-soon cited a 2014 incident where a representative from Kenya, en route to a United Nations biodiversity conference in Pyeongchang, was detained by local immigration officers after accidentally flying to Pyongyang instead.[1]
The official pictograms for 24 sports across 15 disciplines were revealed in January 2017 and are designed using the Korean alphabet as an inspiration.[29]
Mascots[edit]
Main article: Soohorang and Bandabi
Soohorang (수호랑), a white tiger, and Bandabi (반다비), an Asiatic black bear, were announced as the official mascots of the 2018 Winter Olympics and Paralympics.
Sponsors[edit]
[hide]Sponsors of the 2018 Winter Olympics [30]
Worldwide Olympic Partners
Alibaba Group
Atos
Bridgestone
The Coca-Cola Company
DowDuPont (Dow Chemical Company)
General Electric
Intel
Omega SA
Panasonic
Procter & Gamble
Samsung Electronics
Toyota
Visa Inc.
Official Partners
Korean Air
KT Corporation
LG Electronics
POSCO
SK Group (SK Telecom)
VF Corporation (The North Face)
Lotte
McDonald's
Korea Electric Power Corporation
Official Sponsors
EF Education First
Hanwha (Hanwha Life Insurance)
Naver
Samsung Group (Samsung Life Insurance and Samsung Fire & Marine Insurance)
SsangYong Information & Communication Corp
Aggreko
KEB Hana Bank
Shinsegae Food
Official Suppliers
BKL (Bae, Kim and Lee)
Bombardier Inc.
Daedong Industrial
Hanjin Group (Hanjin and Hanjin Travel)
Hanssem
Huawei
Pagoda Academy
PricewaterhouseCoopers (Samil)
S-Oil
Samsung Group (Samsung C&T Corporation and Samsung Securities)
Seoul Tent
Hyundai Department Store
Official Supporters
AhnLab, Inc.
DYFlag
Interpark
Hancom
Douzone
Smart Planet
Daegwanryoung Hanwoo
Ottogi
Noodle Lovers
Cheorwon Odae-ssal
KukDong Metal Fence Co.,Ltd
Influential Inc.
Kelly Services
Somunsa
Maeil Dairies
Heongseong County
DY Corporation
Broadcasting rights[edit]
Main article: List of 2018 Winter Olympics broadcasters
Broadcast rights to the 2018 Winter Olympics in some countries were already sold as part of long-term broadcast rights deals, including the Games' local rightsholder SBS—who extended its rights to the Olympics through 2024 in July 2011.[31]
On 29 June 2015, the IOC announced that Discovery Communications—owner of Eurosport, had acquired exclusive rights to the Olympics from 2018 through 2024 across Europe, excluding Russia, on all platforms. Discovery will sub-license its broadcast rights to local free-to-air networks on a territorial basis. Discovery's rights deal will, initially, not cover France due to pre-existing rights deals with France Télévisions that run through the 2020 Games. Unlike previous pan-European deals, such as with the European Broadcasting Union and Sportfive, Discovery will not solely serve as a reseller, and intends to carry coverage on its regional properties, but has committed to sub-licensing at least 100 hours of coverage to free-to-air networks..[32][33][34] In the United Kingdom, Discovery will hold exclusive pay television rights under license from the BBC, in return for the BBC sub-licensing the free-to-air rights to the 2022 and 2024 Olympics from Discovery.[35]
In the United States, the Games will once again be broadcast by NBC under a long-term contract; it will also be NBC's first Olympics without long-time host Bob Costas, who announced his retirement from the role in favour of Mike Tirico on 7 February 2017.[36][37] On 28 March 2017, NBC also announced that it would air most primetime coverage simultaneously in all time zones, rather than tape-delayed for the west coast.[38]
Concerns and controversies[edit]
Russian Doping[edit]
Russia could possibly face being banned because of their state-sponsored doping program. A decision on the matter will be made in the months prior to the games. [39]
Security[edit]
See also: North Korea and weapons of mass destruction
On 20 September 2017, South Korea’s President Moon Jae-in said the country is pushing to ensure security at Pyeongchang Winter Olympic Games amid rising tensions over the North Korea’s nuclear test and a series of missile launches.[40] However, on the next day, French Minister of Youth Affairs and Sports Laura Flessel-Colovic said France’s Winter Olympics team will not participate in the games in South Korea if its security cannot be guaranteed.[41]
On 22 September 2017, Austria and Germany joined France in staying away from the games. Karl Stoss, head of Austria’s national Olympic committee, said that, "if the situation worsens and the security of our athletes is no longer guaranteed, we will not go to South Korea." The German interior ministry said the security question and the possibility of keeping the German team at home would be addressed “in good time” by the government. Several days later, Laura Flessel-Colovic reaffirmed France's participation in the games. [42]
See also[edit]
Olympics portal
2018 Winter Paralympics
2018 Summer Youth Olympics
Notes and references[edit]
^ Jump up to: a b "Olympics: 2018 Winter Olympics … not in Pyongyang". Manila Bulletin. Agence France-Presse. 26 January 2016. Retrieved 26 January 2016.
Jump up ^ Longman, Jeré; Sang-hun, Choe (2011-07-06). "2018 Winter Games to Be Held in Pyeongchang, South Korea". The New York Times. ISSN 0362-4331. Retrieved 2017-06-07.
Jump up ^ "Pyeongchang 2018 reveal ticket prices for Winter Olympic Games".
Jump up ^ "Gunilla Lindberg to Chair PyeongChang 2018 Coordination Commission". Archived from the original on 18 September 2011. Retrieved 4 October 2011.
Jump up ^ "Coordination Commissions". Olympic.org. Retrieved 8 August 2012.
Jump up ^ "PyeongChang 2018 Organizing Committee Launched". GamesBids.com. Archived from the original on 20 October 2011. Retrieved 8 August 2012.
Jump up ^ "PyeongChang 2018 Praised". Gamesbids.com. Archived from the original on 14 June 2012. Retrieved 8 August 2012.
Jump up ^ "Pyeongchang 2018 have "good grasp of what is expected" says Lindberg after first IOC Coordination Commission visit". Insidethegames.biz. 22 March 2012. Retrieved 8 August 2012.
Jump up ^ "Construction Begins on High-Speed Railway, Critical for PyeongChang 2018 Olympic Games". Gamesbids.com. Archived from the original on 19 June 2012. Retrieved 8 August 2012.
Jump up ^ "IPC Orientates PyeongChang 2018". Gamesbids.com. Archived from the original on 3 August 2012. Retrieved 8 August 2012.
Jump up ^ "Pyeongchang 2018 on "right track" declares Rogge after first visit". Insidethegames.biz. 1 February 2013. Retrieved 19 February 2014.
Jump up ^ "2018평창동계올림픽대회 및 장애인동계올림픽대회 마스코트 아이디어 공모".
Jump up ^ "Pyeongchang 2018 recruits college student reporters: WINNERS". 18 June 2015. Retrieved 5 November 2015.
Jump up ^ "PyeongChang 2018 Alpensia Resort and water park complete and full for summer season". Sportsfeatures.com. Archived from the original on 12 March 2012. Retrieved 8 August 2012.
Jump up ^ "Pyeongchang2018 Volume 2 (Sport and Venues)" (PDF). Archived from the original (PDF) on 18 March 2012. Retrieved 8 August 2012.
^ Jump up to: a b "Pyeongchang 2018 move venue for Opening and Closing Ceremonies | Winter Olympics 2018". insidethegames.biz. 6 July 2012. Retrieved 8 August 2012.
Jump up ^ "Winter Olympics: Big air, mixed curling among new 2018 events". BBC Sport. Retrieved 21 June 2017.
Jump up ^ "NHL will not participate in 2018 Pyeongchang Olympic Games". Sportsnet.ca. Rogers Media. Retrieved 3 April 2017.
Jump up ^ "Olympics and N.H.L. Face Off Over Who Pays to Insure Players". The New York Times. 19 May 2016. Retrieved 21 May 2016.
Jump up ^ "IOC decides not to cover costs for NHL players at Olympics". Sportsnet.ca. Rogers Media. 23 April 2016. Retrieved 21 May 2016.
Jump up ^ Seravalli, Frank. "How the NHL can prevent players from going to Olympics". TSN.ca. Retrieved 4 April 2017.
Jump up ^ "Pyeongchang 2018: Athletes to travel through demilitarised zone". BBC News. May 18, 2017. Retrieved June 8, 2017.
Jump up ^ "With one year until 2018 Winter Games, Russia's status murky". 2017-02-09.
Jump up ^ "Quota allocation for Alpine skiing". www.data.fis-ski.com/. International Ski Federation (FIS). 8 August 2017. Retrieved 8 August 2017.
Jump up ^ "2018 Winter Olympics". IIHF. iihf.com. Retrieved 18 May 2015.
Jump up ^ "Quota allocation for Cross-country skiing". www.data.fis-ski.com/. International Ski Federation (FIS). 8 August 2017. Retrieved 8 August 2017.
Jump up ^ "Pyeongchang 2018 Winter Olympics". worldcurling.org. Retrieved 23 July 2016.
Jump up ^ "PyeongChang 2018 Launches Official Emblem". olympic.org. International Olympic Committee. Retrieved 12 August 2016.
Jump up ^ "PyeongChang 2018 Pictograms". pyeongchang2018.com. January 25, 2017. Retrieved May 30, 2017.
Jump up ^ https://www.pyeongchang2018.com/en/partners
Jump up ^ "IOC awards SBS broadcast rights for 2018, 2020, 2022 and 2024 Olympic Games - Olympic News". International Olympic Committee. 2016-12-09. Retrieved 2017-08-07.
Jump up ^ "IOC awards European broadcast rights to SPORTFIVE". ESPN. 18 February 2009. Retrieved 13 August 2013.
Jump up ^ "Discovery Lands European Olympic Rights Through ’24". Sports Business Journal. Retrieved 1 July 2015.
Jump up ^ "BBC dealt another blow after losing control of TV rights for Olympics". The Guardian. Retrieved 30 June 2015.
Jump up ^ "Olympics coverage to remain on BBC after Discovery deal". The Guardian. Retrieved 7 June 2016.
Jump up ^ "Bob Costas steps down as NBC host of Olympics; Mike Tirico to replace him". USA Today. Retrieved 9 February 2017.
Jump up ^ "Brennan: Bob Costas has been the face of the Olympics for Americans". USA Today. Retrieved 9 February 2017.
Jump up ^ "NBC to Broadcast Winter Olympics Live Across All Time Zones". Variety. Retrieved 28 March 2017.
Jump up ^ "With one year until 2018 Winter Games, Russia's status murky". 2017-02-09.
Jump up ^ "South Korea's Moon says pushing to guarantee safety at Pyeongchang Olympics". Reuters. 20 September 2017.
Jump up ^ "Olympics - France to skip 2018 Winter Games if security not assured". Reuters. 22 September 2017.
Jump up ^ "Olympics: North Korea triggers 2018 Winter Olympics security scare". The Straits Times. 23 September 2017.
External links[edit]
Wikimedia Commons has media related to 2018 Winter Olympics.
Wikivoyage has a travel guide for Pyeongchang 2018 Winter Olympics.
Pyeongchang 2018
Pyeongchang 2018 (IOC)
Preceded by
Sochi Winter Olympics
Pyeongchang
XXIII Olympic Winter Games (2018) Succeeded by
Beijing
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text:
Thylakoid
From Wikipedia, the free encyclopedia
Jump to navigation Jump to search
"Granum" redirects here. For the town in Canada, see Granum, Alberta.
Cell biology
The chloroplast
Components of a typical chloroplast
1 Granum
2 Chloroplast envelope
2.1 Outer membrane
2.2 Intermembrane space
2.3 Inner membrane
3 Thylakoid ◄ You are here
3.1 Thylakoid space (lumen)
3.2 Thylakoid membrane
4 Stromal thylakoid
5 Stroma
6 Nucleoid (DNA ring)
7 Ribosome
8 Plastoglobulus
9 Starch granule

Thylakoids (dark green) inside a chloroplast
A thylakoid is a membrane-bound compartment inside chloroplasts and cyanobacteria. They are the site of the light-dependent reactions of photosynthesis. Thylakoids consist of a thylakoid membrane surrounding a thylakoid lumen. Chloroplast thylakoids frequently form stacks of disks referred to as grana (singular: granum). Grana are connected by intergranal or stroma thylakoids, which join granum stacks together as a single functional compartment.
Contents
1 Etymology
2 Structure
2.1 Membrane
2.2 Lumen
2.3 Granum and stroma lamellae
3 Formation
4 Isolation and fractionation
5 Proteins
5.1 Integral membrane proteins
5.1.1 Photosystems
5.1.2 Cytochrome b6f complex
5.1.3 ATP synthase
5.2 Lumen proteins
5.3 Protein expression
5.4 Protein targeting to the thylakoids
6 Function
6.1 Water photolysis
6.2 Electron transport chains
6.3 Chemiosmosis
6.3.1 Source of proton gradient
6.3.2 ATP generation
7 Thylakoid membranes in cyanobacteria
8 See also
9 References
9.1 Textbook sources
Etymology[edit]
The word thylakoid comes from the Greek word thylakos meaning "sac" or "pouch".[1] Thus, thylakoid means "sac-like" or "pouch-like".
Structure[edit]
Thylakoid structures
TEM image of grana
Thylakoids are membrane-bound structures embedded in the chloroplast stroma. A stack of thylakoids is called a granum and resembles a stack of coins.
Membrane[edit]
The thylakoid membrane is the site of the light-dependent reactions of photosynthesis with the photosynthetic pigments embedded directly in the membrane. It is an alternating pattern of dark and light bands measuring each 1 nanometre.[2] The thylakoid lipid bilayer shares characteristic features with prokaryotic membranes and the inner chloroplast membrane. For example, acidic lipids can be found in thylakoid membranes, cyanobacteria and other photosynthetic bacteria and are involved in the functional integrity of the photosystems.[3] The thylakoid membranes of higher plants are composed primarily of phospholipids[4] and galactolipids that are asymmetrically arranged along and across the membranes.[5] Thylakoid membranes are richer in galactolipids rather than phospholipids; also they predominantly consist of hexagonal phase II forming monogalacotosyl diglyceride lipid. Despite this unique composition, plant thylakoid membranes have been shown to assume largely lipid-bilayer dynamic organization.[6] Lipids forming the thylakoid membranes, richest in high-fluidity linolenic acid[7] are synthesized in a complex pathway involving exchange of lipid precursors between the endoplasmic reticulum and inner membrane of the plastid envelope and transported from the inner membrane to the thylakoids via vesicles.[8]
Lumen[edit]
The thylakoid lumen is a continuous aqueous phase enclosed by the thylakoid membrane. It plays an important role for photophosphorylation during photosynthesis. During the light-dependent reaction, protons are pumped across the thylakoid membrane into the lumen making it acidic down to pH 4.
Granum and stroma lamellae[edit]
In higher plants thylakoids are organized into a granum-stroma membrane assembly. A granum (plural grana) is a stack of thylakoid discs. Chloroplasts can have from 10 to 100 grana. Grana are connected by stroma thylakoids, also called intergranal thylakoids or lamellae. Grana thylakoids and stroma thylakoids can be distinguished by their different protein composition. Grana contribute to chloroplasts' large surface area to volume ratio. Different interpretations of electron tomography imaging of thylakoid membranes has resulted in two models for grana structure. Both posit that lamellae intersect grana stacks in parallel sheets, though whether these sheets intersect in planes perpendicular to the grana stack axis, or are arranged in a right-handed helix is debated.[9]
Formation[edit]
Chloroplasts develop from proplastids when seedlings emerge from the ground. Thylakoid formation requires light. In the plant embryo and in the absence of light, proplastids develop into etioplasts that contain semicrystalline membrane structures called prolamellar bodies. When exposed to light, these prolamellar bodies develop into thylakoids. This does not happen in seedlings grown in the dark, which undergo etiolation. An underexposure to light can cause the thylakoids to fail. This causes the chloroplasts to fail resulting in the death of the plant.
Thylakoid formation requires the action of vesicle-inducing protein in plastids 1 (VIPP1). Plants cannot survive without this protein, and reduced VIPP1 levels lead to slower growth and paler plants with reduced ability to photosynthesize. VIPP1 appears to be required for basic thylakoid membrane formation, but not for the assembly of protein complexes of the thylakoid membrane.[10] It is conserved in all organisms containing thylakoids, including cyanobacteria,[11] green algae, such as Chlamydomonas,[12] and higher plants, such as Arabidopsis thaliana.[13]
Isolation and fractionation[edit]
Thylakoids can be purified from plant cells using a combination of differential and gradient centrifugation.[14] Disruption of isolated thylakoids, for example by mechanical shearing, releases the lumenal fraction. Peripheral and integral membrane fractions can be extracted from the remaining membrane fraction. Treatment with sodium carbonate (Na2CO3) detaches peripheral membrane proteins, whereas treatment with detergents and organic solvents solubilizes integral membrane proteins.
Proteins[edit]
Thylakoid disc with embedded and associated proteins.
Thylakoids contain many integral and peripheral membrane proteins, as well as lumenal proteins. Recent proteomics studies of thylakoid fractions have provided further details on the protein composition of the thylakoids.[15] These data have been summarized in several plastid protein databases that are available online.[16][17]
According to these studies, the thylakoid proteome consists of at least 335 different proteins. Out of these, 89 are in the lumen, 116 are integral membrane proteins, 62 are peripheral proteins on the stroma side, and 68 peripheral proteins on the lumenal side. Additional low-abundance lumenal proteins can be predicted through computational methods.[14][18] Of the thylakoid proteins with known functions, 42% are involved in photosynthesis. The next largest functional groups include proteins involved in protein targeting, processing and folding with 11%, oxidative stress response (9%) and translation (8%).[16]
Integral membrane proteins[edit]
Thylakoid membranes contain integral membrane proteins which play an important role in light harvesting and the light-dependent reactions of photosynthesis. There are four major protein complexes in the thylakoid membrane:
Photosystems I and II
Cytochrome b6f complex
ATP synthase
Photosystem III is located mostly in the grana thylakoids, whereas photosystem I and ATP synthase are mostly located in the stroma thylakoids and the outer layers of grana. The cytochrome b6f complex is distributed evenly throughout thylakoid membranes. Due to the separate location of the two photosystems in the thylakoid membrane system, mobile electron carriers are required to shuttle electrons between them. These carriers are plastoquinone and plastocyanin. Plastoquinone shuttles electrons from photosystem II to the cytochrome b6f complex, whereas plastocyanin carries electrons from the cytochrome b6f complex to photosystem I.
Together, these proteins make use of light energy to drive electron transport chains that generate a chemiosmotic potential across the thylakoid membrane and NADPH, a product of the terminal redox reaction. The ATP synthase uses the chemiosmotic potential to make ATP during photophosphorylation.
Photosystems[edit]
Main article: Photosystem
These photosystems are light-driven redox centers, each consisting of an antenna complex that uses chlorophylls and accessory photosynthetic pigments such as carotenoids and phycobiliproteins to harvest light at a variety of wavelengths. Each antenna complex has between 250 and 400 pigment molecules and the energy they absorb is shuttled by resonance energy transfer to a specialized chlorophyll a at the reaction center of each photosystem. When either of the two chlorophyll a molecules at the reaction center absorbs energy, an electron is excited and transferred to an electron-acceptor molecule. Photosystem I contains a pair of chlorophyll a molecules, designated P700, at its reaction center that maximally absorbs 700 nm light. Photosystem II contains P680 chlorophyll that absorbs 680 nm light best (note that these wavelengths correspond to deep red – see the visible spectrum). The P is short for pigment and the number is the specific absorption peak in nanometers for the chlorophyll molecules in each reaction center.
Cytochrome b6f complex[edit]
Main article: Cytochrome b6f complex
The cytochrome b6f complex is part of the thylakoid electron transport chain and couples electron transfer to the pumping of protons into the thylakoid lumen. Energetically, it is situated between the two photosystems and transfers electrons from photosystem II-plastoquinone to plastocyanin-photosystem I.
ATP synthase[edit]
Main article: ATP synthase
The thylakoid ATP synthase is a CF1FO-ATP synthase similar to the mitochondrial ATPase. It is integrated into the thylakoid membrane with the CF1-part sticking into stroma. Thus, ATP synthesis occurs on the stromal side of the thylakoids where the ATP is needed for the light-independent reactions of photosynthesis.
Lumen proteins[edit]
The electron transport protein plastocyanin is present in the lumen and shuttles electrons from the cytochrome b6f protein complex to photosystem I. While plastoquinones are lipid-soluble and therefore move within the thylakoid membrane, plastocyanin moves through the thylakoid lumen.
The lumen of the thylakoids is also the site of water oxidation by the oxygen evolving complex associated with the lumenal side of photosystem II.
Lumenal proteins can be predicted computationally based on their targeting signals. In Arabidopsis, out of the predicted lumenal proteins possessing the Tat signal, the largest groups with known functions are 19% involved in protein processing (proteolysis and folding), 18% in photosynthesis, 11% in metabolism, and 7% redox carriers and defense.[14]
Protein expression[edit]
Chloroplasts have their own genome, which encodes a number of thylakoid proteins. However, during the course of plastid evolution from their cyanobacterial endosymbiotic ancestors, extensive gene transfer from the chloroplast genome to the cell nucleus took place. This results in the four major thylakoid protein complexes being encoded in part by the chloroplast genome and in part by the nuclear genome. Plants have developed several mechanisms to co-regulate the expression of the different subunits encoded in the two different organelles to assure the proper stoichiometry and assembly of these protein complexes. For example, transcription of nuclear genes encoding parts of the photosynthetic apparatus is regulated by light. Biogenesis, stability and turnover of thylakoid protein complexes are regulated by phosphorylation via redox-sensitive kinases in the thylakoid membranes.[19] The translation rate of chloroplast-encoded proteins is controlled by the presence or absence of assembly partners (control by epistasy of synthesis).[20] This mechanism involves negative feedback through binding of excess protein to the 5' untranslated region of the chloroplast mRNA.[21] Chloroplasts also need to balance the ratios of photosystem I and II for the electron transfer chain. The redox state of the electron carrier plastoquinone in the thylakoid membrane directly affects the transcription of chloroplast genes encoding proteins of the reaction centers of the photosystems, thus counteracting imbalances in the electron transfer chain.[22]
Protein targeting to the thylakoids[edit]
Schematic representation of thylakoid protein targeting pathways.[23]
Thylakoid proteins are targeted to their destination via signal peptides and prokaryotic-type secretory pathways inside the chloroplast. Most thylakoid proteins encoded by a plant's nuclear genome need two targeting signals for proper localization: An N-terminal chloroplast targeting peptide (shown in yellow in the figure), followed by a thylakoid targeting peptide (shown in blue). Proteins are imported through the translocon of outer and inner membrane (Toc and Tic) complexes. After entering the chloroplast, the first targeting peptide is cleaved off by a protease processing imported proteins. This unmasks the second targeting signal and the protein is exported from the stroma into the thylakoid in a second targeting step. This second step requires the action of protein translocation components of the thylakoids and is energy-dependent. Proteins are inserted into the membrane via the SRP-dependent pathway (1), the Tat-dependent pathway (2), or spontaneously via their transmembrane domains (not shown in figure). Lumenal proteins are exported across the thylakoid membrane into the lumen by either the Tat-dependent pathway (2) or the Sec-dependent pathway (3) and released by cleavage from the thylakoid targeting signal. The different pathways utilize different signals and energy sources. The Sec (secretory) pathway requires ATP as energy source and consists of SecA, which binds to the imported protein and a Sec membrane complex to shuttle the protein across. Proteins with a twin arginine motif in their thylakoid signal peptide are shuttled through the Tat (twin arginine translocation) pathway, which requires a membrane-bound Tat complex and the pH gradient as an energy source. Some other proteins are inserted into the membrane via the SRP (signal recognition particle) pathway. The chloroplast SRP can interact with its target proteins either post-translationally or co-translationally, thus transporting imported proteins as well as those that are translated inside the chloroplast. The SRP pathway requires GTP and the pH gradient as energy sources. Some transmembrane proteins may also spontaneously insert into the membrane from the stromal side without energy requirement.[23]
Function[edit]
Light-dependent reactions of photosynthesis at the thylakoid membrane
The thylakoids are the site of the light-dependent reactions of photosynthesis. These include light-driven water oxidation and oxygen evolution, the pumping of protons across the thylakoid membranes coupled with the electron transport chain of the photosystems and cytochrome complex, and ATP synthesis by the ATP synthase utilizing the generated proton gradient.
Water photolysis[edit]
Main article: photolysis
The first step in photosynthesis is the light-driven reduction (splitting) of water to provide the electrons for the photosynthetic electron transport chains as well as protons for the establishment of a proton gradient. The water-splitting reaction occurs on the lumenal side of the thylakoid membrane and is driven by the light energy captured by the photosystems. This oxidation of water conveniently produces the waste product O2 that is vital for cellular respiration. The molecular oxygen formed by the reaction is released into the atmosphere.
Electron transport chains[edit]
Main article: electron transport chain
Two different variations of electron transport are used during photosynthesis:
Noncyclic electron transport or Non-cyclic photophosphorylation produces NADPH + H+ and ATP.
Cyclic electron transport or Cyclic photophosphorylation produces only ATP.
The noncyclic variety involves the participation of both photosystems, while the cyclic electron flow is dependent on only photosystem I.
Photosystem I uses light energy to reduce NADP+ to NADPH + H+, and is active in both noncyclic and cyclic electron transport. In cyclic mode, the energized electron is passed down a chain that ultimately returns it (in its base state) to the chlorophyll that energized it.
Photosystem II uses light energy to oxidize water molecules, producing electrons (e−), protons (H+), and molecular oxygen (O2), and is only active in noncyclic transport. Electrons in this system are not conserved, but are rather continually entering from oxidized 2H2O (O2 + 4 H+ + 4 e−) and exiting with NADP+ when it is finally reduced to NADPH.
Chemiosmosis[edit]
Main article: chemiosmosis
A major function of the thylakoid membrane and its integral photosystems is the establishment of chemiosmotic potential. The carriers in the electron transport chain use some of the electron's energy to actively transport protons from the stroma to the lumen. During photosynthesis, the lumen becomes acidic, as low as pH 4, compared to pH 8 in the stroma.[24] This represents a 10,000 fold concentration gradient for protons across the thylakoid membrane.
Source of proton gradient[edit]
The protons in the lumen come from three primary sources.
Photolysis by photosystem II oxidises water to oxygen, protons and electrons in the lumen.
The transfer of electrons from photosystem II to plastoquinone during non-cyclic electron transport consumes two protons from the stroma. These are released in the lumen when the reduced plastoquinol is oxidized by the cytochrome b6f protein complex on the lumen side of the thylakoid membrane. From the plastoquinone pool, electrons pass through the cytochrome b6f complex. This integral membrane assembly resembles cytochrome bc1.
The reduction of plastoquinone by ferredoxin during cyclic electron transport also transfers two protons from the stroma to the lumen.
The proton gradient is also caused by the consumptions of protons in the stroma to make NADPH from NADP+ at the NADP reductase.
ATP generation[edit]
The molecular mechanism of ATP (Adenosine triphosphate) generation in chloroplasts is similar to that in mitochondria and takes the required energy from the proton motive force (PMF).[citation needed] However, chloroplasts rely more on the chemical potential of the PMF to generate the potential energy required for ATP synthesis. The PMF is the sum of a proton chemical potential (given by the proton concentration gradient) and a transmembrane electrical potential (given by charge separation across the membrane). Compared to the inner membranes of mitochondria, which have a significantly higher membrane potential due to charge separation, thylakoid membranes lack a charge gradient.[citation needed] To compensate for this, the 10,000 fold proton concentration gradient across the thylakoid membrane is much higher compared to a 10 fold gradient across the inner membrane of mitochondria. The resulting chemiosmotic potential between the lumen and stroma is high enough to drive ATP synthesis using the ATP synthase. As the protons travel back down the gradient through channels in ATP synthase, ADP + Pi are combined into ATP. In this manner, the light-dependent reactions are coupled to the synthesis of ATP via the proton gradient.[citation needed]
Thylakoid membranes in cyanobacteria[edit]
Thylakoids (green) inside a cyanobacterium (Synechocystis)
Cyanobacteria are photosynthetic prokaryotes with highly differentiated membrane systems. Cyanobacteria have an internal system of thylakoid membranes where the fully functional electron transfer chains of photosynthesis and respiration reside. The presence of different membrane systems lends these cells a unique complexity among bacteria. Cyanobacteria must be able to reorganize the membranes, synthesize new membrane lipids, and properly target proteins to the correct membrane system. The outer membrane, plasma membrane, and thylakoid membranes each have specialized roles in the cyanobacterial cell. Understanding the organization, functionality, protein composition and dynamics of the membrane systems remains a great challenge in cyanobacterial cell biology.[25]
The thylakoid membranes of the cyanobacteria are not differentiated into granum and stroma regions as observed in plants. They form stacks of parallel sheets close to the cytoplasmic membrane with a low packing density.[26] The relatively large distance between the thylakoids provides space for the external light harvesting antennae, the phycobilisomes.[27] This macrostructure, as in the case of higher plants, shows some flexibility during changes in the physicochemical environment.[28]
See also[edit]
Arthur Meyer (botanist)
André Jagendorf
Chemiosmosis
Electrochemical gradient
Endosymbiosis
Oxygen evolution
Photosynthesis
References[edit]
Jump up ^ θύλακος. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
Jump up ^ "Photosynthesis" McGraw Hill Encyclopedia of Science and Technology, 10th ed. 2007. Vol. 13 p. 469
Jump up ^ Sato N (2004). "Roles of the acidic lipids sulfoquinovosyl diacylglycerol and phosphatidylglycerol in photosynthesis: their specificity and evolution". J Plant Res. 117 (6): 495–505. doi:10.1007/s10265-004-0183-1. PMID 15538651.
Jump up ^ "photosynthesis."Encyclopædia Britannica. 2008. Encyclopædia Britannica 2006 Ultimate Reference Suite DVD 9 Apr. 2008
Jump up ^ Spraque SG (1987). "Structural and functional organization of galactolipids on thylakoid membrane organization". J Bioenerg Biomembr. 19 (6): 691–703. doi:10.1007/BF00762303. PMID 3320041.
Jump up ^ YashRoy, R.C. (1990). "Magnetic resonance studies of dynamic organization of lipids in chloroplast membranes" (PDF). Journal of Biosciences. 15 (4): 281–288. doi:10.1007/bf02702669.
Jump up ^ YashRoy, R.C. (1987). "13C NMR studies of lipid fatty-acyl chains of chloroplast membranes". Indian Journal of Biochemistry and Biophysics. 24: 177–178.
Jump up ^ Benning C, Xu C, Awai K (2006). "Non-vesicular and vesicular lipid trafficking involving plastids". Curr Opin Plant Biol. 9 (3): 241–7. doi:10.1016/j.pbi.2006.03.012. PMID 16603410.
Jump up ^ Mustárdy, L.; Buttle, K.; Steinbach, G.; Garab, G. (2008). "The Three-Dimensional Network of the Thylakoid Membranes in Plants: Quasihelical Model of the Granum-Stroma Assembly". Plant Cell. 20 (10): 2552–2557. doi:10.1105/tpc.108.059147. PMC 2590735. PMID 18952780.
Jump up ^ Elena Aseeva; Friederich Ossenbühl; Claudia Sippel; Won K. Cho; Bernhard Stein; Lutz A. Eichacker; Jörg Meurer; Gerhard Wanner; Peter Westhoff; Jürgen Soll; Ute C. Vothknecht (2007). "Vipp1 is required for basic thylakoid membrane formation but not for the assembly of thylakoid protein complexes". Plant Physiol Biochem. 45 (2): 119–28. doi:10.1016/j.plaphy.2007.01.005. PMID 17346982.
Jump up ^ Westphal S, Heins L, Soll J, Vothknecht U (2001). "Vipp1 deletion mutant of Synechocystis: A connection between bacterial phage shock and thylakoid biogenesis?". Proc Natl Acad Sci USA. 98 (7): 4243–8. doi:10.1073/pnas.061501198. PMC 31210. PMID 11274448.
Jump up ^ Liu C, Willmund F, Golecki J, Cacace S, Markert C, Heß B, Schroda M, Schroda M (2007). "The chloroplast HSP70B-CDJ2-CGE1 chaperones catalyse assembly and disassembly of VIPP1 oligomers in Chlamydomonas". Plant J. 50 (2): 265–77. doi:10.1111/j.1365-313X.2007.03047.x. PMID 17355436.
Jump up ^ Kroll D, Meierhoff K, Bechtold N, Kinoshita M, Westphal S, Vothknecht U, Soll J, Westhoff P (2001). "VIPP1, a nuclear gene of Arabidopsis thaliana essential for thylakoid membrane formation". Proc Natl Acad Sci USA. 98 (7): 4238–42. doi:10.1073/pnas.061500998. PMC 31209. PMID 11274447.
^ Jump up to: a b c Peltier J, Emanuelsson O, Kalume D, Ytterberg J, Friso G, Rudella A, Liberles D, Söderberg L, Roepstorff P, von Heijne G, van Wijk KJ (2002). "Central Functions of the Lumenal and Peripheral Thylakoid Proteome of Arabidopsis Determined by Experimentation and Genome-Wide Prediction". Plant Cell. 14 (1): 211–36. doi:10.1105/tpc.010304. PMC 150561. PMID 11826309.
Jump up ^ van Wijk K (2004). "Plastid proteomics". Plant Physiol Biochem. 42 (12): 963–77. doi:10.1016/j.plaphy.2004.10.015. PMID 15707834.
^ Jump up to: a b Friso G, Giacomelli L, Ytterberg A, Peltier J, Rudella A, Sun Q, Wijk K (2004). "In-Depth Analysis of the Thylakoid Membrane Proteome of Arabidopsis thaliana Chloroplasts: New Proteins, New Functions, and a Plastid Proteome Database". Plant Cell. 16 (2): 478–99. doi:10.1105/tpc.017814. PMC 341918. PMID 14729914. - The Plastid Proteome Database
Jump up ^ Kleffmann T, Hirsch-Hoffmann M, Gruissem W, Baginsky S (2006). "plprot: a comprehensive proteome database for different plastid types". Plant Cell Physiol. 47 (3): 432–6. doi:10.1093/pcp/pcj005. PMID 16418230. – Plastid Protein Database
Jump up ^ Peltier J, Friso G, Kalume D, Roepstorff P, Nilsson F, Adamska I, van Wijk K (2000). "Proteomics of the Chloroplast: Systematic Identification and Targeting Analysis of Lumenal and Peripheral Thylakoid Proteins". Plant Cell. 12 (3): 319–41. doi:10.1105/tpc.12.3.319. PMC 139834. PMID 10715320.
Jump up ^ Vener AV, Ohad I, Andersson B (1998). "Protein phosphorylation and redox sensing in chloroplast thylakoids". Curr Opin Plant Biol. 1 (3): 217–23. doi:10.1016/S1369-5266(98)80107-6. PMID 10066592.
Jump up ^ Choquet Y, Wostrikoff K, Rimbault B, Zito F, Girard-Bascou J, Drapier D, Wollman F (2001). "Assembly-controlled regulation of chloroplast gene translation". Biochem Soc Trans. 29 (Pt 4): 421–6. doi:10.1042/BST0290421. PMID 11498001.
Jump up ^ Minai L, Wostrikoff K, Wollman F, Choquet Y (2006). "Chloroplast Biogenesis of Photosystem II Cores Involves a Series of Assembly-Controlled Steps That Regulate Translation". Plant Cell. 18 (1): 159–75. doi:10.1105/tpc.105.037705. PMC 1323491. PMID 16339851.
Jump up ^ Allen J, Pfannschmidt T (2000). "Balancing the two photosystems: photosynthetic electron transfer governs transcription of reaction centre genes in chloroplasts". Philos Trans R Soc Lond B Biol Sci. 355 (1402): 1351–9. doi:10.1098/rstb.2000.0697. PMC 1692884. PMID 11127990.
^ Jump up to: a b Gutensohn M, Fan E, Frielingsdorf S, Hanner P, Hou B, Hust B, Klösgen R (2006). "Toc, Tic, Tat et al.: structure and function of protein transport machineries in chloroplasts". J. Plant Physiol. 163 (3): 333–47. doi:10.1016/j.jplph.2005.11.009. PMID 16386331.
Jump up ^ Jagendorf A. T. and E. Uribe (1966). "ATP formation caused by acid-base transition of spinach chloroplasts". Proc. Natl. Acad. Sci. USA. 55: 170–177. doi:10.1073/pnas.55.1.170. PMC 285771.
Jump up ^ Herrero A and Flores E (editor). (2008). The Cyanobacteria: Molecular Biology, Genomics and Evolution (1st ed.). Caister Academic Press. ISBN 1-904455-15-8. [1].
Jump up ^ van de Meene, AM; Hohmann-Marriott, MF; Vermaas, WF; Roberson, RW (January 2006). "The three-dimensional structure of the cyanobacterium Synechocystis sp. PCC 6803". Archives of Microbiology. 184 (5): 259–70. doi:10.1007/s00203-005-0027-y. PMID 16320037.
Jump up ^ Olive, J; Ajlani, G; Astier, C; Recouvreur, M; Vernotte, C. "Ultrastructure and light adaptation of phycobilisome mutants of Synechocystis PCC 6803". Biochimica et Biophysica Acta (BBA) - Bioenergetics. 1319 (2–3): 275–282. doi:10.1016/S0005-2728(96)00168-5.
Jump up ^ Nagy, G; Posselt, D; Kovács, L; Holm, JK; Szabó, M; Ughy, B; Rosta, L; Peters, J; Timmins, P; Garab, G (Jun 1, 2011). "Reversible membrane reorganizations during photosynthesis in vivo: revealed by small-angle neutron scattering". The Biochemical Journal. 436 (2): 225–30. doi:10.1042/BJ20110180. PMID 21473741.
Textbook sources[edit]
Heller, H. Craig; Orians, Gordan H.; Purves, William K. & Sadava, David (2004). LIFE: The Science of Biology (7th ed.). Sinauer Associates, Inc. ISBN 0-7167-9856-5.
Raven, Peter H.; Ray F. Evert; Susan E. Eichhorn (2005). Biology of Plants (7th ed.). New York: W.H. Freeman and Company Publishers. pp. 115–127. ISBN 0-7167-1007-2.
Herrero A and Flores E (editors). (2008). The Cyanobacteria: Molecular Biology, Genomics and Evolution (1st ed.). Caister Academic Press. ISBN 978-1-904455-15-8.
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https://sh.wikipedia.org/wiki/Tilakoid
Tilakoid – Serbo-Croatian
sh
sh
interlanguage-link-target

values:
interlanguage-link interwiki-sh
display: none;

values:
https://fi.wikipedia.org/wiki/Tylakoidi
Tylakoidi – Finnish
fi
fi
interlanguage-link-target

values:
interlanguage-link interwiki-fi
display: none;

values:
https://sv.wikipedia.org/wiki/Tylakoid
Tylakoid – Swedish
sv
sv
interlanguage-link-target

values:
interlanguage-link interwiki-sv
display: none;

values:
https://tr.wikipedia.org/wiki/Tilakoid
Tilakoid – Turkish
tr
tr
interlanguage-link-target

values:
interlanguage-link interwiki-tr
display: none;

values: Тилакоїд – Ukrainian

values:
interlanguage-link interwiki-uk
display: none;

values: All languages (initial selection from common choices by you and others)

values: mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject page-Thylakoid rootpage-Thylakoid skin-vector action-view