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Information Technology Act, 2000
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Information Technology Act, 2000
the Act to provide legal recognition for transactions carried out by means of electronic data interchange and other means of electronic communication, commonly referred to as "electronic commerce", which involve the use of alternatives to paper-based methods of communication and storage of information, to nusta editing electronic filing of documents with the Government agencies and further to amend the Indian Penal Code, the Indian Evidence Act, 1872, the Bankers' Books Evidence Act, 1891 and the Reserve Bank of India Act, 1934 and for matters connected therewith or incidental thereto.
Citation
Information Technology Act, 2000
Enacted by
Parliament of India
Date enacted
9 June 2000
Date assented to
9 June 2000
Date signed
9 May 2000
Date commenced
17 October 2000
Amendments
IT (Amendment) Act 2008
Status: In force
The Information Technology Act, 2000 (also known as ITA-2000, or the IT Act) is an Act of the Indian Parliament (No 21 of 2000) notified on 17 October 2000. It is the primary law in India dealing with cybercrime and electronic commerce. It is based on the United Nations Model Law on Electronic Commerce 1996 (UNCITRAL Model) recommended by the General Assembly of United Nations by a resolution dated 30 January 1997.[1]
Contents
[hide]
1 Background
2 Summary
2.1 Amendments
2.2 Offences
3 Notable cases
3.1 Section 66
3.2 Section 66A
4 Criticisms
4.1 Section 66A and restriction of free speech
4.1.1 Resolution for amendment
4.1.2 Petitions challenging constitutionality
4.1.3 Revocation by the Supreme Court
4.2 Strict data privacy rules
4.3 Mandatory decryption
5 Future changes
6 See also
7 References
8 Further reading
9 External links
Background[edit]
The bill was passed in the budget session of 2000 and signed by President K. R. Narayanan on 9 May 2000. The bill was finalised by group of officials headed by then Minister of Information Technology Pramod Mahajan.[2]
Summary[edit]
The original Act contained 94 sections, divided in 13 chapters and 4 schedules. The laws apply to the whole of India. Persons of other nationalities can also be indicted under the law, if the crime involves a computer or network located in India.[3]
The Act provides legal framework for electronic governance by giving recognition to electronic records and digital signatures. The formations of Controller of Certifying Authorities was directed by the Act, to regulate issuing of digital signatures. It also defines cyber crimes and prescribed penalties for them. It also established a Cyber Appellate Tribunal to resolve disputes rising from this new law.[3] The Act also amended various sections of Indian Penal Code, 1860, Indian Evidence Act, 1872, Banker's Book Evidence Act, 1891, and Reserve Bank of India Act, 1934 to make them compliant with new technologies.[3]
Amendments[edit]
A major amendment was made in 2008. It introduced the Section 66A which penalized sending of "offensive messages". It also introduced the Section 69, which gave authorities the power of "interception or monitoring or decryption of any information through any computer resource". It also introduced for child porn, cyber terrorism and voyeurism. It was passed on 22 December 2008 without any debate in Lok Sabha. The next day it was passed by the Rajya Sabha. It was signed by the then President (Pratibha Patil) on 5 February 2009.[4][5][6][7]
Offences[edit]
List of offences and the corresponding penalties:[8][9]
Section
Offence
Description
Penalty
65 Tampering with computer source documents If a person knowingly or intentionally conceals, destroys or alters or intentionally or knowingly causes another to conceal, destroy or alter any computer source code used for a computer, computer programme, computer system or computer network, when the computer source code is required to be kept or maintained by law for the time being in force. Imprisonment up to three years, or/and with fine up to ₹200,000
66 Hacking with computer system If a person with the intent to cause or knowing that he is likely to cause wrongful loss or damage to the public or any person destroys or deletes or alters any information residing in a computer resource or diminishes its value or utility or affects it injuriously by any means, commits hack. Imprisonment up to three years, or/and with fine up to ₹500,000
66B Receiving stolen computer or communication device A person receives or retains a computer resource or communication device which is known to be stolen or the person has reason to believe is stolen. Imprisonment up to three years, or/and with fine up to ₹100,000
66C Using password of another person A person fradulently uses the password, digital signature or other unique identification of another person. Imprisonment up to three years, or/and with fine up to ₹100,000
66D Cheating using computer resource If a person cheats someone using a computer resource or communication. Imprisonment up to three years, or/and with fine up to ₹100,000
66E Publishing private images of others If a person captures, transmits or publishes images of a person's private parts without his/her consent or knowledge. Imprisonment up to three years, or/and with fine up to ₹200,000
66F Acts of cyberterrorism If a person denies access to an authorised personnel to a computer resource, accesses a protected system or introduces contaminant into a system, with the intention of threatening the unity, integrity, sovereignty or security of India, then he commits cyberterrorism. Imprisonment up to life.
67 Publishing information which is obscene in electronic form. If a person publishes or transmits or causes to be published in the electronic form, any material which is lascivious or appeals to the prurient interest or if its effect is such as to tend to deprave and corrupt persons who are likely, having regard to all relevant circumstances, to read, see or hear the matter contained or embodied in it. Imprisonment up to five years, or/and with fine up to ₹1,000,000
67A Publishing images containing sexual acts If a person publishes or transmits images containing a sexual explicit act or conduct. Imprisonment up to seven years, or/and with fine up to ₹1,000,000
67B Publishing child porn or predating children online If a person captures, publishes or transmits images of a child in a sexually explicit act or conduct. If a person induces a child into a sexual act. A child is defined as anyone under 18. Imprisonment up to five years, or/and with fine up to ₹1,000,000 on first conviction. Imprisonment up to seven years, or/and with fine up to ₹1,000,000 on second conviction.
67C Failure to maintain records Persons deemed as intermediatary (such as an ISP) must maintain required records for stipulated time. Failure is an offence. Imprisonment up to three years, or/and with fine.
68 Failure/refusal to comply with orders The Controller may, by order, direct a Certifying Authority or any employee of such Authority to take such measures or cease carrying on such activities as specified in the order if those are necessary to ensure compliance with the provisions of this Act, rules or any regulations made thereunder. Any person who fails to comply with any such order shall be guilty of an offence. Imprisonment up to three years, or/and with fine up to {{currency}} – invalid amount (help)
69 Failure/refusal to decrypt data If the Controller is satisfied that it is necessary or expedient so to do in the interest of the sovereignty or integrity of India, the security of the State, friendly relations with foreign Stales or public order or for preventing incitement to the commission of any cognizable offence, for reasons to be recorded in writing, by order, direct any agency of the Government to intercept any information transmitted through any computer resource. The subscriber or any person in charge of the computer resource shall, when called upon by any agency which has been directed, must extend all facilities and technical assistance to decrypt the information. The subscriber or any person who fails to assist the agency referred is deemed to have committed a crime. Imprisonment up to seven years and possible fine.
70 Securing access or attempting to secure access to a protected system The appropriate Government may, by notification in the Official Gazette, declare that any computer, computer system or computer network to be a protected system.
The appropriate Government may, by order in writing, authorise the persons who are authorised to access protected systems. If a person who secures access or attempts to secure access to a protected system, then he is committing an offence.
Imprisonment up to ten years, or/and with fine.
71 Misrepresentation If anyone makes any misrepresentation to, or suppresses any material fact from, the Controller or the Certifying Authority for obtaining any license or Digital Signature Certificate. Imprisonment up to three years, or/and with fine up to ₹100,000
Notable cases[edit]
Section 66[edit]
In February 2001, in one of the first cases, the Delhi police arrested two men running a web-hosting company. The company had shut down a website over non-payment of dues. The owner of the site had claimed that he had already paid and complained to the police. The Delhi police had charged the men for hacking under Section 66 of the IT Act and breach of trust under Section 408 of the Indian Penal Code. The two men had to spend 6 days in Tihar jail waiting for bail. Bhavin Turakhia, chief executive officer of directi.com, a webhosting firm said that this interpretation of the law would be problematic for web-hosting companies.[10]
In February 2017, M/s Voucha Gram India Pvt.Ltd, owner of Delhi based Ecommerce Portal www.gyftr.com made a Complaint with Hauz Khas Police Station against some hackers from different cities accusing them for IT Act / Theft / Cheating / Misappropriation / Criminal Conspiracy / Criminal Breach of Trust / Cyber Crime of Hacking / Snooping / Tampering with Computer source documents and the Web Site and extending the threats of dire consequences to employees, as a result four hackers were arrested by South Delhi Police for Digital Shoplifting.[11]
Section 66A[edit]
In September 2012, a freelance cartoonist Aseem Trivedi was arrested under the Section 66A of the IT Act, Section 2 of Prevention of Insults to National Honour Act, 1971 and for sedition under the Section 124 of the Indian Penal Code.[12] His cartoons depicting widespread corruption in India were considered offensive.[13][14]
On 12 April 2012, a Chemistry professor from Jadavpur University, Ambikesh Mahapatra, was arrested for sharing a cartoon of West Bengal Chief Minister Mamata Banerjee and then Railway Minister Mukul Roy.[15] The email was sent from the email address of a housing society. Subrata Sengupta, the secretary of the housing society, was also arrested. They were charged under Section 66A and B of the IT Act, for defamation under Sections 500, for obscene gesture to a woman under Section 509, and abetting a crime under Section 114 of the Indian Penal Code.[16]
On 30 October 2012, a Puducherry businessman Ravi Srinivasan was arrested under Section 66A. He had sent tweet accusing Karti Chidambaram, son of then Finance Minister P. Chidambaram, of corruption. Karti Chidambaram had complained to the police.[17]
On 19 November 2012, a 21-year-old girl was arrested from Palghar for posting a message on Facebook criticising the shutdown in Mumbai for the funeral of Bal Thackeray. Another 20-year-old girl was arrested for "liking" the post. They were initially charged under Section 295A of the Indian Penal Code (hurting religious sentiments) and Section 66A of the IT Act. Later, Section 295A was replaced by Section 505(2) (promoting enmity between classes).[18] A group of Shiv Sena workers vandalised a hospital run by the uncle of one of girls.[19] On 31 January 2013, a local court dropped all charges against the girls.[20]
On 18 March 2015, a teenaged boy was arrested from Bareilly, Uttar Pradesh, for making a post on Facebook insulting politician Azam Khan. The post allegedly contained hate speech against a community and was falsely attributed to Azam Khan by the boy. He was charged under Section 66A of the IT Act, and Sections 153A (promoting enmity between different religions), 504 (intentional insult with intent to provoke breach of peace) and 505 (public mischief) of Indian Penal Code. After the Section 66A was repealed on 24 March, the state government said that they would continue the prosecution under the remaining charges.[21][22]
Criticisms[edit]
Section 66A and restriction of free speech[edit]
From its establishment as an amendment to the original act in 2008, Section 66A attracted controversy over its unconstitutional nature:
Section
Offence
Description
Penalty
66A Publishing offensive, false or threatening information Any person who sends by any means of a computer resource any information that is grossly offensive or has a menacing character; or any information which he knows to be false, but for the purpose of causing annoyance, inconvenience, danger, obstruction, insult shall be punishable with imprisonment for a term which may extend to three years and with fine. Imprisonment up to three years, with fine.
Resolution for amendment[edit]
In December 2012, P Rajeev, a Rajya Sabha member from Kerala, tried to pass a resolution seeking to amend the Section 66A. He was supported by D. Bandyopadhyay, Gyan Prakash Pilania, Basavaraj Patil Sedam, Narendra Kumar Kashyap, Rama Chandra Khuntia and Baishnab Charan Parida. P Rajeev pointed that cartoons and editorials allowed in traditional media, were being censored in the new media. He also said that law was barely debated before being passed in December 2008.[23]
Rajeev Chandrasekhar suggested the 66A should only apply to person to person communication pointing to a similar section under the Indian Post Office Act, 1898. Shantaram Naik opposed any changes, saying that the misuse of law was sufficient to warrant changes. Then Minister for Communications and Information Technology Kapil Sibal defended the existing law, saying that similar laws existed in US and UK. He also said that a similar provision existed under Indian Post Office Act, 1898. However, P Rajeev said that the UK dealt only with communication from person to person.[23]
Petitions challenging constitutionality[edit]
In November 2012, IPS officer Amitabh Thakur and his wife social activist Nutan Thakur, filed a petition in the Lucknow bench of the Allahabad High Court claiming that the Section 66A violated the freedom of speech guaranteed in the Article 19(1)(a) of the Constitution of India. They said that the section was vague and frequently misused.[24]
Also in November 2012, a Delhi-based law student, Shreya Singhal, filed a Public Interest Litigation (PIL) in the Supreme Court of India. She argued that the Section 66A was vaguely phrased, as result it violated Article 14, 19 (1)(a) and Article 21 of the Constitution. The PIL was accepted on 29 November 2012.[25][26] A similar petition was also filed by the founder of MouthShut.com, Faisal Farooqui,[27] and NGO Common Cause represented by Prashant Bhushan[28] In August 2014, the Supreme Court asked the central government to respond to petitions filed by Mouthshut.com and later petition filed by the Internet and Mobile Association of India (IAMAI) which claimed that the IT Act gave the government power to arbitrarily remove user-generated content.[29]
Revocation by the Supreme Court[edit]
On 24 March 2015, the Supreme Court of India, gave the verdict that Section 66A is unconstitutional in entirety. The court said that Section 66A of IT Act 2000 is "arbitrarily, excessively and disproportionately invades the right of free speech" provided under Article 19(1) of the Constitution of India. But the Court turned down a plea to strike down sections 69A and 79 of the Act, which deal with the procedure and safeguards for blocking certain websites. [30][31]
Strict data privacy rules[edit]
The data privacy rules introduced in the Act in 2011 have been described as too strict by some Indian and US firms. The rules require firms to obtain written permission from customers before collecting and using their personal data. This has affected US firms which outsource to Indian companies. However, some companies have welcomed the strict rules, saying it will remove fears of outsourcing to Indian companies.[32]
Mandatory decryption[edit]
See also: Mandatory decryption
The Section 69 allows intercepting any information and ask for information decryption. To refuse decryption is an offence. The Indian Telegraph Act, 1885 allows the government to tap phones. But, according to a 1996 Supreme Court verdict the government can tap phones only in case of a "public emergency". But, there is no such restriction on Section 69.[5]
Future changes[edit]
On 2 April 2015, the Chief Minister of Maharashtra, Devendra Fadnavis revealed to the state assembly that a new law was being framed to replace the repealed Section 66A. Fadnavis was replying to a query Shiv Sena leader Neelam Gorhe. Gorhe had said that repeal of the law would encourage online miscreants and asked whether the state government would frame a law to this regard. Fadnavis said that the previous law had resulted in no convictions, so the law would be framed such that it would be strong and result in convictions.[33]
On 13 April 2015, it announced that the Ministry of Home Affairs would form a committee of officials from the Intelligence Bureau, Central Bureau of Investigation, National Investigation Agency, Delhi Police and ministry itself to produce a new legal framework. This step was reportedly taken after complaints from intelligence agencies that, they were no longer able to counter online posts that involved national security matter or incite people to commit an offence, such as online recruitment for ISIS.[34][35] Former Minister of State with the Ministry of Information Technology, Milind Deora has supported a new "unambiguous section to replace 66A".[36]
See also[edit]
Chilling effect
Mouthshut.com v. Union of India
Save Your Voice
References[edit]
Jump up ^ B.M.Gandhi. Indian Penal Code. India: Eastern Book Company. p. 41. ISBN 9788170128922.
Jump up ^ "IT Act to come into force from August 15". Rediff. 9 August 2000. Retrieved 14 April 2015.
^ Jump up to: a b c Sujata Pawar; Yogesh Kolekar (23 March 2015). Essentials of Information Technology Law. Notion Press. pp. 296–306. ISBN 978-93-84878-57-3. Retrieved 14 April 2015.
Jump up ^ "Section 66A of the Information Technology Act". Centre for Internet and Society (India). Retrieved 14 April 2015.
^ Jump up to: a b "Yes, snooping's allowed". The Indian Express. 6 February 2009. Retrieved 14 April 2015.
Jump up ^ "Deaf, Dumb & Dangerous - 21 Minutes: That was the time our MPs spent on Section 66A. How they played". The Telegraph (India). 26 March 2015. Retrieved 6 May 2015.
Jump up ^ "Amended IT Act to prevent cyber crime comes into effect". The Hindu. 27 October 2015. Retrieved 8 May 2015.
Jump up ^ "The Information Technology (Amendment) Act, 2008". Retrieved 7 May 2017.
Jump up ^ Chapter 11: Offences
Jump up ^ "Cyber crime that wasn't?". Rediff. 19 February 2001. Retrieved 14 April 2015.
Jump up ^ "Fourt Hackers Arrested in Delhi, Cyber Crime, Gift Vouchers, Hacking, Section 65 / 66 of IT Act, Gyftr". Information Technology Act. 2010-02-10. Retrieved 2017-05-07.
Jump up ^ "'If Speaking The Truth Is Sedition, Then I Am Guilty'". Outlook India. 10 September 2010. Retrieved 14 April 2015.
Jump up ^ "Indian cartoonist Aseem Trivedi jailed after arrest on sedition charges". The Guardian. 10 September 2010. Retrieved 14 April 2015.
Jump up ^ Section 66A: Punishment for sending offensive messages through communication service, etc.
Jump up ^ "Professor arrested for poking fun at Mamata". Hindustan Times. 14 April 2012. Retrieved 14 April 2015.
Jump up ^ "Cartoon a conspiracy, prof an offender: Mamata". Hindustan Times. 13 April 2012. Retrieved 14 April 2015.
Jump up ^ "Arrest over tweet against Chidambaram's son propels 'mango man' Ravi Srinivasan into limelight". India Today. 2 November 2012. Retrieved 14 April 2015.
Jump up ^ "Mumbai shuts down due to fear, not respect". The Hindu. 19 November 2012. Retrieved 23 April 2015.
Jump up ^ "FB post: 10 Sainiks arrested for hospital attack". The Hindu. 20 November 2012. Retrieved 23 April 2015.
Jump up ^ "Facebook row: Court scraps charges against Palghar girls". The Hindu. 31 January 2013. Retrieved 23 April 2015.
Jump up ^ "Teen arrested for Facebook post attributed to Azam Khan gets bail". The Times of India. 19 March 2015. Retrieved 6 May 2015.
Jump up ^ "UP tells SC that prosecution on boy for post against Azam Khan will continue". The Indian Express. 24 April 2015. Retrieved 6 May 2015.
^ Jump up to: a b "Section 66A of IT Act undemocratic: RS MPs". The Times of India. 15 December 2012. Retrieved 6 May 2015.
Jump up ^ "After Mumbai FB case, writ filed in Lucknow to declare section 66A, IT Act 2000 as ultra-vires". The Times of India. 21 November 2012. Retrieved 14 April 2015.
Jump up ^ "SC accepts PIL challenging Section 66A of IT Act". The Times of India. 29 November 2012. Retrieved 23 April 2015.
Jump up ^ "Shreya Singhal: The student who took on India's internet laws". BBC News. 24 March 2015. Retrieved 6 May 2015.
Jump up ^ "'Heavens Won't Fall' if Controversial Parts of IT Act are Stayed, Says Supreme Court". NDTV. 4 December 2014. Retrieved 6 May 2015.
Jump up ^ http://www.newslaundry.com/2015/03/24/the-heroes-in-the-battle-against-section-66a/
Jump up ^ "SC seeks govt reply on PIL challenging powers of IT Act". Live Mint. 30 August 2015. Retrieved 6 May 2015.
Jump up ^ "SC strikes down 'draconian' Section 66A". The Hindu. 25 March 2015. Retrieved 23 April 2015.
Jump up ^ "SC quashes Section 66A of IT Act: Key points of court verdict". The Times of India. 24 March 2015. Retrieved 6 May 2015.
Jump up ^ "India data privacy rules may be too strict for some U.S. companies". The Washington Post. 21 May 2011. Retrieved 23 April 2015.
Jump up ^ "Centre working on new law similar to Section 66A: Devendra Fadnavis". The Times of India. 2 April 2015. Retrieved 6 May 2015.
Jump up ^ "Section 66A of the IT Act likely to be back in softer avatar". The Economic Times. 14 April 2015. Retrieved 6 May 2015.
Jump up ^ "New panel to work on Section 66A alternative". Hindustan Times. 14 April 2015. Retrieved 6 May 2015.
Jump up ^ "Former IT minister Milind Deora: Why we need a new Section 66A". Rediff. 2 April 2015. Retrieved 6 May 2015.
Further reading[edit]
Shreya Singhal v. Union of India (Supreme Court of India 24 March 2015). Text
David Rizk (10 June 2011). "New Indian Internet Intermediary Regulations Pose Serious Threats to Net Users' Freedom of Expression". Electronic Frontier Foundation.
External links[edit]
"Information Technology Act, 2000".
"Information Technology Act, 2008".
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Nucleolus
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This article may be too technical for most readers to understand. Please help improve it to make it understandable to non-experts, without removing the technical details. The talk page may contain suggestions. (October 2012) (Learn how and when to remove this template message)
The nucleolus is contained within the cell nucleus.
Images from "The Nucleolus"[1] DIC images
tripartite organization
granular component
The nucleolus (/njuːˈkliːələs/ or /ˌnjuːkliˈoʊləs/, plural nucleoli /njuːˈkliːəˌlaɪ/ or /ˌnjuːkliˈoʊlaɪ/) is the largest structure in the nucleus of eukaryotic cells.[2] It is best known as the site of ribosome biogenesis. Nucleoli also participate in the formation of signal recognition particles and play a role in the cell's response to stress.[3] Nucleoli are made of proteins, DNA and RNA and form around specific chromosomal regions called nucleolar organizing regions. Malfunction of nucleoli can be the cause of several human conditions called "nucleolopathies"[4] and the nucleolus is being investigated as a target for cancer chemotherapy.[5][6]
Contents
[hide]
1 History
2 Structure
3 Function and ribosome assembly
4 Sequestration of proteins
5 See also
6 References
7 External links
History[edit]
The nucleolus was identified by bright-field microscopy during the 1830s.[1] Little was known about the function of the nucleolus until 1964, when a study[7] of nucleoli by John Gurdon and Donald Brown in the African clawed frog Xenopus laevis generated increasing interest in the function and detailed structure of the nucleolus. They found that 25% of the frog eggs had no nucleolus and that such eggs were not capable of life. Half of the eggs had one nucleolus and 25% had two. They concluded that the nucleolus had a function necessary for life. In 1966 Max L. Birnstiel and collaborators showed via nucleic acid hybridization experiments that DNA within nucleoli code for ribosomal RNA.[8][9]
Transcription in the FC, fibrillarin & nucleophosmin[10]
Structure[edit]
Three major components of the nucleolus are recognized: the fibrillar center (FC), the dense fibrillar component (DFC), and the granular component (GC).[2] Transcription of the rDNA occurs in the FC.[10] The DFC contains the protein fibrillarin,[10] which is important in rRNA processing. The GC contains the protein nucleophosmin,[10] (B23 in the external image) which is also involved in ribosome biogenesis.
Structure of the eukaryotic rDNA repeat.[2]
However, it has been proposed that this particular organization is only observed in higher eukaryotes and that it evolved from a bipartite organization with the transition from anamniotes to amniotes. Reflecting the substantial increase in the DNA intergenic region, an original fibrillar component would have separated into the FC and the DFC.[11]
Play media
Nucleus from a cell line. Fibrillarin in red. Transcription regulatory protein CTCFL in green. Nuclear DNA in blue.
Another structure identified within many nucleoli (particularly in plants) is a clear area in the center of the structure referred to as a nucleolar vacuole.[12] Nucleoli of various plant species have been shown to have very high concentrations of iron[13] in contrast to human and animal cell nucleoli.
The nucleolus ultrastructure can be visualized through an electron microscope, while the organization and dynamics can be studied through fluorescent protein tagging and fluorescent recovery after photobleaching (FRAP). Antibodies against the PAF49 protein can also be used as a marker for the nucleolus in immunofluorescence experiments.[14]
Function and ribosome assembly[edit]
Electron micrograph of part of a HeLa cell. The image is a screen capture from this movie, which shows a Z-stack of the cell.
In ribosome biogenesis, two of the three eukaryotic RNA polymerases (pol I and III) are required, and these function in a coordinated manner. In an initial stage, the rRNA genes are transcribed as a single unit within the nucleolus by RNA polymerase I. In order for this transcription to occur, several pol I-associated factors and DNA-specific trans-acting factors are required. In yeast, the most important are: UAF (upstream activating factor), TBP (TATA-box binding protein), and CF (core factor), which bind promoter elements and form the preinitiation complex (PIC), which is in turn recognized by RNA pol. In humans, a similar PIC is assembled with SL1, the promoter selectivity factor (composed of TBP and TBP-associated factors, or TAFs), transcription initiation factors, and UBF (upstream binding factor). RNA polymerase I transcribes most rRNA transcripts (28S, 18S, and 5.8S) but the 5S rRNA subunit (component of the 60S ribosomal subunit) is transcribed by RNA polymerase III.[15]
Transcription of rRNA yields a long precursor molecule (45S pre-rRNA) which still contains the ITS and ETS. Further processing is needed to generate the 18S RNA, 5.8S and 28S RNA molecules. In eukaryotes, the RNA-modifying enzymes are brought to their respective recognition sites by interaction with guide RNAs, which bind these specific sequences. These guide RNAs belong to the class of small nucleolar RNAs (snoRNAs) which are complexed with proteins and exist as small-nucleolar-ribonucleoproteins (snoRNPs). Once the rRNA subunits are processed, they are ready to be assembled into larger ribosomal subunits. However, an additional rRNA molecule, the 5S rRNA, is also necessary. In yeast, the 5S rDNA sequence is localized in the intergenic spacer and is transcribed in the nucleolus by RNA pol.
In higher eukaryotes and plants, the situation is more complex, for the 5S DNA sequence lies outside the Nucleolus Organiser Region (NOR) and is transcribed by RNA pol III in the nucleoplasm, after which it finds its way into the nucleolus to participate in the ribosome assembly. This assembly not only involves the rRNA, but ribosomal proteins as well. The genes encoding these r-proteins are transcribed by pol II in the nucleoplasm by a "conventional" pathway of protein synthesis (transcription, pre-mRNA processing, nuclear export of mature mRNA and translation on cytoplasmic ribosomes). The mature r-proteins are then "imported" back into the nucleus and finally the nucleolus. Association and maturation of rRNA and r-proteins result in the formation of the 40S (small) and 60S (large) subunits of the complete ribosome. These are exported through the nuclear pore complexes to the cytoplasm, where they remain free or become associated with the endoplasmic reticulum, forming rough endoplasmic reticulum (RER).[16][17]
In human endometrial cells, a network of nucleolar channels is sometimes formed. The origin and function of this network has not yet been clearly identified.[18]
Sequestration of proteins[edit]
In addition to its role in ribosomal biogenesis, the nucleolus is known to capture and immobilize proteins, a process known as nucleolar detention. Proteins that are detained in the nucleolus are unable to diffuse and to interact with their binding partners. Targets of this post-translational regulatory mechanism include VHL, PML, MDM2, POLD1, RelA, HAND1 and hTERT, among many others. It is now known that long noncoding RNAs originating from intergenic regions of the nucleolus are responsible for this phenomenon.[19]
Cell biology
The animal cell
Components of a typical animal cell:
Nucleolus
Nucleus
Ribosome (little dots)
Vesicle
Rough endoplasmic reticulum
Golgi apparatus (or "Golgi body")
Cytoskeleton
Smooth endoplasmic reticulum
Mitochondrion
Vacuole
Cytosol (fluid that contains organelles, comprising the cytoplasm)
Lysosome
Centrosome
Cell membrane
See also[edit]
Differential interference contrast microscopy
References[edit]
^ Jump up to: a b Pederson T (2011). "The nucleolus". Cold Spring Harbor Perspectives in Biology. 3 (3). PMC 3039934  . PMID 21106648. doi:10.1101/cshperspect.a000638.
^ Jump up to: a b c O'Sullivan JM, Pai DA, Cridge AG, Engelke DR, Ganley AR (2013). "The nucleolus: a raft adrift in the nuclear sea or the keystone in nuclear structure?". Biomolecular Concepts. 4 (3): 277–86. PMC 5100006  . PMID 25436580. doi:10.1515/bmc-2012-0043.
Jump up ^ Olsen, Mark OJ (December 2010). "Nucleolus: Structure and Function". ELS. ISBN 0470016175. doi:10.1002/9780470015902.a0005975.pub2. Retrieved 17 October 2014.
Jump up ^ Hetman M (2014). "Role of the nucleolus in human diseases. Preface". Biochimica et Biophysica Acta. 1842 (6): 757. PMID 24631655. doi:10.1016/j.bbadis.2014.03.004.
Jump up ^ Quin JE, Devlin JR, Cameron D, Hannan KM, Pearson RB, Hannan RD (2014). "Targeting the nucleolus for cancer intervention". Biochimica et Biophysica Acta. 1842 (6): 802–16. PMID 24389329. doi:10.1016/j.bbadis.2013.12.009.
Jump up ^ Woods SJ, Hannan KM, Pearson RB, Hannan RD (2015). "The nucleolus as a fundamental regulator of the p53 response and a new target for cancer therapy". Biochimica et Biophysica Acta. 1849 (7): 821–9. PMID 25464032. doi:10.1016/j.bbagrm.2014.10.007.
Jump up ^ BROWN DD, GURDON JB (1964). "ABSENCE OF RIBOSOMAL RNA SYNTHESIS IN THE ANUCLEOLATE MUTANT OF XENOPUS LAEVIS" (PDF). Proceedings of the National Academy of Sciences of the United States of America. 51: 139–46. PMC 300879  . PMID 14106673.
Jump up ^ Birnstiel ML, Wallace H, Sirlin JL, Fischberg M (1966). "Localization of the ribosomal DNA complements in the nucleolar organizer region of Xenopus laevis". National Cancer Institute Monograph. 23: 431–47. PMID 5963987.
Jump up ^ Wallace H, Birnstiel ML (1966). "Ribosomal cistrons and the nucleolar organizer". Biochimica et Biophysica Acta. 114 (2): 296–310. PMID 5943882.
^ Jump up to: a b c d Sirri V, Urcuqui-Inchima S, Roussel P, Hernandez-Verdun D (2008). "Nucleolus: the fascinating nuclear body". Histochemistry and Cell Biology. 129 (1): 13–31. PMC 2137947  . PMID 18046571. doi:10.1007/s00418-007-0359-6.
Jump up ^ Thiry M, Lafontaine DL (2005). "Birth of a nucleolus: the evolution of nucleolar compartments". Trends Cell Biol. 15 (4): 194–9. PMID 15817375. doi:10.1016/j.tcb.2005.02.007. as PDF
Jump up ^ Beven AF, Lee R, Razaz M, Leader DJ, Brown JW, Shaw PJ (1 June 1996). "The organization of ribosomal RNA processing correlates with the distribution of nucleolar snRNAs". J. Cell. Sci. 109 (6): 1241–51. PMID 8799814.
Jump up ^ Roschzttardtz H, Grillet L, Isaure MP, et al. (2011). "Plant cell nucleolus as a hot spot for iron". J. Biol. Chem. 286 (32): 27863–6. PMC 3151030  . PMID 21719700. doi:10.1074/jbc.C111.269720.
Jump up ^ PAF49 antibody | GeneTex Inc. Genetex.com. Retrieved on 2013-03-03.
Jump up ^ Champe, Pamela C.; Harvey, Richard A.; Ferrier, Denise R. (2005). Lippincott's Illustrated Reviews: Biochemistry. Lippincott Williams & Wilkins. ISBN 978-0-7817-2265-0.
Jump up ^ Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter (2002). Molecular Biology of the Cell (4th ed.). New York: Garland Science. pp. 331–3. ISBN 0-8153-3218-1.
Jump up ^ Cooper, Geoffrey M.; Hausman, Robert E. (2007). The Cell: A Molecular Approach (4th ed.). Sinauer Associates. pp. 371–9. ISBN 0878932208.
Jump up ^ Wang T, Schneider J; Origin and fate of the nucleolar channel system of normal human endometrium; Cell Res. (1992); 2(2), 97-102
Jump up ^ Audas TE, Jacob MD, Lee S (2012). "Immobilization of proteins in the nucleolus by ribosomal intergenic spacer noncoding RNA". Mol Cell. 45 (2): 147–57. PMID 22284675. doi:10.1016/j.molcel.2011.12.012.
External links[edit]
Wikimedia Commons has media related to Cell nucleolus.
Look up nucleolus in Wiktionary, the free dictionary.
Cooper, Geoffrey M. (2000). "The Nucleolus". The Cell: A Molecular Approach (2nd ed.). Sunderland MA: Sinauer Associates. ISBN 0-87893-106-6.
Nucleolus under electron microscope II at uni-mainz.de
Nuclear Protein Database – search under compartment
Cell Nucleolus at the US National Library of Medicine Medical Subject Headings (MeSH)
Histology image: 20104loa – Histology Learning System at Boston University
[show]
v
t
e
Structures of the cell / organelles (TH H1.00.01.2-3)
Endomembrane system
Cell membrane
Nucleus
Nucleolus
Endoplasmic reticulum
Golgi apparatus
Parenthesome
Autophagosome
Vesicles
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Endosome
Phagosome
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[show]
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t
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nuclear lamina
Pore complex:
Nucleoporin
NUP35
NUP37
NUP43
NUP50
NUP54
NUP62
NUP85
NUP88
NUP93
NUP98
NUP107
NUP133
NUP153
NUP155
NUP160
NUP188
NUP205
NUP210
NUP214
AAAS
Nucleolus
Cajal (coiled) body
SMN
GEMIN2
GEMIN4
GEMIN5
GEMIN6
GEMIN7
DDX20
COIL
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PTBP1
CUGBP1
TCOF
ATXN7
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Dot (PML body)
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SMC6
Transition nuclear protein:
TNP1
TNP2
Nuclear matrix (Nucleoskeleton)
Nucleoplasm (Nucleosol)
LITAF
see also transcription factors and intracellular receptors
see also nucleus diseases
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