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Acts of Union 1707
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"Union of England and Scotland" redirects here. It is not to be confused with Union of England and Scotland Act 1603 or Treaty of Union.
Union with Scotland Act 1706[1]
Act of Parliament
Parliament of England
Long title
An Act for a Union of the Two Kingdoms of England and Scotland
Citation
1706 c. 11
Territorial extent
Kingdom of England (inc. Wales); subsequently, United Kingdom
Status: Current legislation
Revised text of statute as amended
Union with England Act 1707
Act of Parliament
Parliament of Scotland
Long title
Act Ratifying and Approving the Treaty of Union of the Two Kingdoms of Scotland and England
Citation
1707 c. 7
Territorial extent
Kingdom of Scotland; subsequently, United Kingdom
Status: Current legislation
Revised text of statute as amended
Constitutional documents relevant to the status of the United Kingdom and legislative unions of its constituent countries
Treaty of Union
1706
Acts of Union
1707
Personal Union of 1714
1714
Wales and Berwick Act
1746
Irish Constitution
1782
Acts of Union
1800
Government of Ireland Act
1920
Anglo-Irish Treaty
1921
Royal and Parliamentary Titles Act
1927
N. Ireland (Temporary Provisions) Act
1972
European Communities Act
1972
Local Government Act
1972
Local Government (Scotland) Act
1973
Northern Ireland Assembly
1973
N. Ireland Constitution Act
1973
Referendum Act
1975
Scotland Act
1978
Wales Act
1978
Local Government (Wales) Act
1994
Local Government etc. (Scotland) Act
1994
Referendums (Scotland & Wales) Act
1997
Good Friday Agreement
1998
Northern Ireland Act
1998
Government of Wales Act
1998
Scotland Act
1998
Government of Wales Act
2006
Northern Ireland Act
2009
European Union Act
2011
Scotland Act
2012
Edinburgh Agreement
2012
Wales Act
2014
European Union Referendum Act
2015
Scotland Act
2016
Wales Act
2017
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The Acts of Union were two Acts of Parliament: the Union with Scotland Act 1706 passed by the Parliament of England, and the Union with England Act passed in 1707 by the Parliament of Scotland. They put into effect the terms of the Treaty of Union that had been agreed on 22 July 1706, following negotiation between commissioners representing the parliaments of the two countries. By the two Acts, the Kingdom of England and the Kingdom of Scotland—which at the time were separate states with separate legislatures, but with the same monarch—were, in the words of the Treaty, "United into One Kingdom by the Name of Great Britain".[2]
The two countries had shared a monarch since the Union of the Crowns in 1603, when King James VI of Scotland inherited the English throne from his double first cousin twice removed, Queen Elizabeth I. Although described as a Union of Crowns, until 1707 there were in fact two separate Crowns resting on the same head (as opposed to the implied creation of a single Crown and a single Kingdom, exemplified by the later Kingdom of Great Britain). There had been three attempts in 1606, 1667, and 1689 to unite the two countries by Acts of Parliament, but it was not until the early 18th century that both political establishments came to support the idea, albeit for different reasons.
The Acts took effect on 1 May 1707. On this date, the Scottish Parliament and the English Parliament united to form the Parliament of Great Britain, based in the Palace of Westminster in London, the home of the English Parliament.[3] Hence, the Acts are referred to as the Union of the Parliaments. On the Union, the historian Simon Schama said "What began as a hostile merger, would end in a full partnership in the most powerful going concern in the world ... it was one of the most astonishing transformations in European history."[4]
Contents
[hide]
1 Historical background
1.1 Previous attempts at union
1.1.1 Early Stuart union
1.1.2 Union during the interregnum
1.1.3 Later attempts
1.2 Treaty and passage of the Acts of 1707
2 Political motivations
2.1 English perspective
2.2 Scottish perspective
3 Provisions of the Acts
4 Evaluations
5 300th anniversary
6 Scottish voting records
7 See also
8 Notes
9 Further reading
10 Other books
11 External links
Historical background[edit]
Previous attempts at union[edit]
England and Scotland were separate states for several centuries before eventual union, and English attempts to take over Scotland by military force in the late 13th and early 14th centuries were ultimately unsuccessful (see the Wars of Scottish Independence). The first attempts at Union surrounded the foreseen unification[clarification needed] of the Royal lines of Scotland and England. In pursuing the English throne in the 1560s, Mary, Queen of Scots pledged herself to a peaceful union between the two kingdoms.[5]
England and Scotland were ruled by the same king for the first time in 1603 when James VI of Scotland also became the king of England. However they remained two separate states until 1 May 1707.
Early Stuart union[edit]
The first Union flag, created by James VI and I, symbolising the uniting of England and Scotland under one Crown (A separate version was used in Scotland during the 17th Century)
The first attempt to unite the parliaments of England and Scotland was by Mary's son, King James VI and I. On his accession to the English throne in 1603 King James announced his intention to unite his two realms so that he would not be "guilty of bigamy". James used his royal prerogative powers to take the style of "King of Great Britain"[6] and to give an explicitly British character to his court and person.[7] Whilst James assumed the creation of a full union was a foregone conclusion, the Parliament of England was concerned that the formation of a new state would deprive England of its ancient liberties, taking on the more absolutist monarchical structure James had previously enjoyed in Scotland.[8] In the meantime, James declared that Great Britain be viewed "as presently united, and as one realm and kingdom, and the subjects of both realms as one people".[9]
The Scottish and English parliaments established a commission to negotiate a union, formulating an instrument of union between the two countries. However, the idea of political union was unpopular, and when James dropped his policy of a speedy union, the topic quietly disappeared from the legislative agenda. When the House of Commons attempted to revive the proposal in 1610, it was met with a more open hostility.[10]
Union during the interregnum[edit]
Main article: Scotland under the Commonwealth
Rare Commonwealth-era flag depicting the union between England and Scotland
The Solemn League and Covenant 1643 sought a forced union of the Church of England into the Church of Scotland, and although the covenant referred repeatedly to union between the three kingdoms of England, Scotland, and Ireland, a political union was not spelled out.
In the aftermath of the Civil War, in which the Covenanters had fought for the King, Oliver Cromwell occupied Scotland and began a process of creating a "Godly Britannic" Union between the former Kingdoms.[11] In 1651, the Parliament of England issued the Tender of Union declaration supporting Scotland's incorporation into the Commonwealth and sent Commissioners to Scotland with the express purpose of securing support for Union, which was assented to by the Commissioners (Members of Parliament) in Scotland. On 12 April 1654, Cromwell – styling himself Lord Protector of England, Scotland and Ireland – enacted An Ordinance by the Protector for the Union of England and Scotland, which created "one Commonwealth and under one Government" to be known as the Commonwealth of England, Scotland and Ireland.[12] The ordinance was ratified by the Second Protectorate Parliament, as an Act of Union, on 26 June 1657.[13] One united Parliament sat in Westminster, with 30 representatives from Scotland and 30 from Ireland joining the existing members from England. Whilst free trade was brought about amongst the new Commonwealth, the economic benefits were generally not felt as a result of heavy taxation used to fund Cromwell's New Model Army.[11]
This republican union was dissolved automatically with the restoration of King Charles II to the thrones of England and Scotland. Scottish members expelled from the Commonwealth Parliament petitioned unsuccessfully for a continuance of the union. Cromwell's union had simultaneously raised interest in and suspicion of the concept of union and when Charles II attempted to recreate the union and fulfil the work of his grandfather in 1669, negotiations between Commissioners ground to a halt.[14]
Later attempts[edit]
An abortive scheme for union occurred in Scotland in 1670.[15]
Following the Glorious Revolution of 1688, the records of the Parliament of Scotland show much discussion of possible union. William III and Mary II, whilst supportive of the idea, had no interest in allowing it to delay their enthronement. Impetus for this incorporating union came almost entirely from King William, who feared leaving Scotland open to a French invasion. In the 1690s, the economic position of Scotland worsened, and relations between Scotland and England became strained.[16] In the following decade, however, union again became a significant topic of political debate.
Treaty and passage of the Acts of 1707[edit]
"Articles of Union otherwise known as Treaty of Union", 1707
Deeper political integration had been a key policy of Queen Anne from the time she acceded to the throne in 1702. Under the aegis of the Queen and her ministers in both kingdoms, the parliaments of England and Scotland agreed to participate in fresh negotiations for a union treaty in 1705.
Both countries appointed 31 commissioners to conduct the negotiations. Most of the Scottish commissioners favoured union, and about half were government ministers and other officials. At the head of the list was Queensberry, and the Lord Chancellor of Scotland, the Earl of Seafield.[17] The English commissioners included the Lord High Treasurer, the Earl of Godolphin, the Lord Keeper, Baron Cowper, and a large number of Whigs who supported union. Tories were not in favour of union and only one was represented among the commissioners.[17]
Negotiations between the English and Scottish commissioners took place between 16 April and 22 July 1706 at the Cockpit in London. Each side had its own particular concerns. Within a few days, England gained a guarantee that the Hanoverian dynasty would succeed Queen Anne to the Scottish crown, and Scotland received a guarantee of access to colonial markets, in the hope that they would be placed on an equal footing in terms of trade.[18]
After negotiations ended in July 1706, the acts had to be ratified by both Parliaments. In Scotland, about 100 of the 227 members of the Parliament of Scotland were supportive of the Court Party. For extra votes the pro-court side could rely on about 25 members of the Squadrone Volante, led by the Marquess of Montrose and the Duke of Roxburghe. Opponents of the court were generally known as the Country party, and included various factions and individuals such as the Duke of Hamilton, Lord Belhaven and Andrew Fletcher of Saltoun, who spoke forcefully and passionately against the union. The Court party enjoyed significant funding from England and the Treasury and included many who had accumulated debts following the Darien Disaster.[19]
In Scotland, the Duke of Queensberry was largely responsible for the successful passage of the Union act by the Scottish Parliament. In Scotland, he received much criticism from local residents, but in England he was cheered for his action. He had received around half of the funding awarded by the Westminster treasury for himself. In April 1707, he travelled to London to attend celebrations at the royal court, and was greeted by groups of noblemen and gentry lined along the road. From Barnet, the route was lined with crowds of cheering people, and once he reached London a huge crowd had formed. On 17 April, the Duke was gratefully received by the Queen at Kensington Palace.[20]
Political motivations[edit]
Portrait of Queen Anne in 1702, the year she became queen, from the school of John Closterman
English perspective[edit]
The English purpose was to ensure that Scotland would not choose a monarch different from the one on the English throne. The two countries had shared a king for much of the previous century, but the English were concerned that an independent Scotland with a different king, even if he were a Protestant, might make alliances against England. The English succession was provided for by the English Act of Settlement 1701, which ensured that the monarch of England would be a Protestant member of the House of Hanover. Until the Union of Parliaments, the Scottish throne might be inherited by a different successor after Queen Anne: the Scottish Act of Security 1704 granted parliament the right to choose a successor and explicitly required a choice different from the English monarch unless the English were to grant free trade and navigation. Many people in England were unhappy about the prospect, however. English overseas possessions made England very wealthy in comparison to Scotland, a poor country with few roads, very little industry and almost no Navy.[citation needed] This made some view unification as a markedly unequal relationship.
Scottish perspective[edit]
In Scotland, some claimed that union would enable Scotland to recover from the financial disaster wrought by the Darien scheme through English assistance and the lifting of measures put in place through the Alien Act to force the Scottish Parliament into compliance with the Act of Settlement.[21]
The combined votes of the Court party with a majority of the Squadrone Volante were sufficient to ensure the final passage of the treaty through the House.
Personal financial interests were also allegedly involved. Many Commissioners had invested heavily in the Darien scheme and they believed that they would receive compensation for their losses; Article 15 granted £398,085 10s sterling to Scotland, a sum known as The Equivalent, to offset future liability towards the English national debt. In essence it was also used as a means of compensation for investors in the Company of Scotland's Darien scheme, as 58.6% was allocated to its shareholders and creditors.[22]
18thC French illustration of an opening of the Scottish Parliament
Even more direct bribery was also said to be a factor.[23] £20,000 (£240,000 Scots) was dispatched to Scotland for distribution by the Earl of Glasgow. James Douglas, 2nd Duke of Queensberry, the Queen's Commissioner in Parliament, received £12,325, more than 60% of the funding. (Some contend that all of this money was properly accounted for as compensation for loss of office, pensions and so forth not outwith the usual run of government. It is perhaps a debate that will never be set to rest. However, modern research has shown that payments were made to supporters of union that appear not to have been overdue salaries. At least four payments were made to people who were not even members of the Scottish Parliament.) Robert Burns referred to this:
We're bought and sold for English Gold,
Such a Parcel of Rogues in a Nation.
Some of the money was used to hire spies, such as Daniel Defoe; his first reports were of vivid descriptions of violent demonstrations against the Union. "A Scots rabble is the worst of its kind," he reported, "for every Scot in favour there is 99 against". Years later, Sir John Clerk of Penicuik, originally a leading Unionist, wrote in his memoirs that Defoe "was a spy among us, but not known as such, otherwise the Mob of Edinburgh would pull him to pieces."
The Treaty could be considered very unpopular at the time. Popular unrest occurred in Edinburgh, as mentioned above, with some lesser but still substantial riots in Glasgow. The people of Edinburgh demonstrated against the treaty, and their apparent leader in opposition to the Unionists, was James Hamilton, 4th Duke of Hamilton. However, Hamilton was actually on the side of the English Government. Demonstrators in Edinburgh were opposed to the Union, for many reasons: they feared the Kirk would be Anglicised; that Anglicisation would remove democracy from the only really elementally democratic part of the Kingdom and they feared that tax rises would come.[24]
Sir George Lockhart of Carnwath, the only member of the Scottish negotiating team against union, noted that "The whole nation appears against the Union"[25] and even Sir John Clerk of Penicuik, an ardent pro-unionist and Union negotiator, observed that the treaty was "contrary to the inclinations of at least three-fourths of the Kingdom".[25] Public opinion against the Treaty as it passed through the Scottish Parliament was voiced through petitions from shires, burghs, presbyteries and parishes. The Convention of Royal Burghs also petitioned against the Union as proposed:
That it is our indispensable duty to signify to your grace that, as we are not against an honourable and safe union with England far less can we expect to have the condition of the people of Scotland, with relation to these great concerns, made better and improved without a Scots Parliament.[26]
Not one petition in favour of an incorporating union was received by Parliament. On the day the treaty was signed, the carilloner in St Giles Cathedral, Edinburgh, rang the bells in the tune Why should I be so sad on my wedding day?[27] Threats of widespread civil unrest resulted in Parliament imposing martial law.
Provisions of the Acts[edit]
Royal heraldic badge of Queen Anne, depicting the Tudor rose and the Scottish thistle growing out of the same stem.
Main article: Treaty of Union
The Treaty of Union, agreed between representatives of the Parliament of England and the Parliament of Scotland in 1706, consisted of 25 articles, 15 of which were economic in nature. In Scotland, each article was voted on separately and several clauses in articles were delegated to specialised subcommittees. Article 1 of the treaty was based on the political principle of an incorporating union and this was secured by a majority of 116 votes to 83 on 4 November 1706. To minimise the opposition of the Church of Scotland, an Act was also passed to secure the Presbyterian establishment of the Church, after which the Church stopped its open opposition, although hostility remained at lower levels of the clergy. The treaty as a whole was finally ratified on 16 January 1707 by a majority of 110 votes to 69.[28]
The two Acts incorporated provisions for Scotland to send representative peers from the Peerage of Scotland to sit in the House of Lords. It guaranteed that the Church of Scotland would remain the established church in Scotland, that the Court of Session would "remain in all time coming within Scotland", and that Scots law would "remain in the same force as before". Other provisions included the restatement of the Act of Settlement 1701 and the ban on Roman Catholics from taking the throne. It also created a customs union and monetary union.
The Act provided that any "laws and statutes" that were "contrary to or inconsistent with the terms" of the Act would "cease and become void."
Soon after the Union, the Act 6 Anne c.40—later infelicitously named the Union with Scotland (Amendment) Act 1707—united the English and Scottish Privy Councils and decentralised Scottish administration by appointing justices of the peace in each shire to carry out administration. In effect it took the day-to-day government of Scotland out of the hands of politicians and into those of the College of Justice.
Evaluations[edit]
Scotland benefited, says historian G.N. Clark, gaining "freedom of trade with England and the colonies" as well as "a great expansion of markets." The agreement guaranteed the permanent status of the Presbyterian church in Scotland, and the separate system of laws and courts in Scotland. Clark argued that in exchange for the financial benefits and bribes that England bestowed, what it gained was:
of inestimable value. Scotland accepted the Hanoverian succession and gave up her power of threatening England's military security and complicating her commercial relations....The sweeping successes of the eighteenth-century wars owed much to the new unity of the two nations.[29]
By the time Samuel Johnson and James Boswell made their tour in 1773, recorded in A Journey to the Western Islands of Scotland, Johnson noted that Scotland was "a nation of which the commerce is hourly extending, and the wealth increasing" and in particular that Glasgow had become one of the greatest cities of Britain.[30]
300th anniversary[edit]
The £2 coin issued in the United Kingdom in 2007 to commemorate the 300th anniversary of the Acts of Union
A commemorative two-pound coin was issued to mark the tercentennial—300th anniversary—of the Union, which occurred two days before the Scottish Parliament general election on 3 May 2007.[31]
The Scottish Executive held a number of commemorative events through the year including an education project led by the Royal Commission on the Ancient and Historical Monuments of Scotland, an exhibition of Union-related objects and documents at the National Museums of Scotland and an exhibition of portraits of people associated with the Union at the National Galleries of Scotland.[32]
Scottish voting records[edit]
Map of commissioner voting on the ratification of the Treaty of Union.
Voting Records for 16 January 1707 ratification of the Treaty of Union
Commissioner
Constituency/Position
Party
Vote
James Graham, 1st Duke of Montrose Lord President of the Council of Scotland/Stirlingshire Court Party Yes
John Campbell, 2nd Duke of Argyll Court Party Yes
John Hay, 2nd Marquess of Tweeddale Squadrone Volante Yes
William Kerr, 2nd Marquess of Lothian Court Party Yes
John Erskine, 22nd Earl of Mar Court Party Yes
John Gordon, 16th Earl of Sutherland Court Party Yes
John Hamilton-Leslie, 9th Earl of Rothes Squadrone Volante Yes
James Douglas, 11th Earl of Morton Yes
William Cunningham, 12th Earl of Glencairn Yes
James Hamilton, 6th Earl of Abercorn Yes
John Ker, 1st Duke of Roxburghe Squadrone Volante Yes
Thomas Hamilton, 6th Earl of Haddington Yes
John Maitland, 5th Earl of Lauderdale Yes
David Wemyss, 4th Earl of Wemyss Yes
William Ramsay, 5th Earl of Dalhousie Yes
James Ogilvy, 4th Earl of Findlater Banffshire Yes
David Leslie, 3rd Earl of Leven Yes
David Carnegie, 4th Earl of Northesk Yes
Earl of Belcarras Yes
Archibald Douglas, 1st Earl of Forfar Yes
William Boyd, 3rd Earl of Kilmarnock Yes
John Keith, 1st Earl of Kintore Yes
Patrick Hume, 1st Earl of Marchmont Squadrone Volante Yes
George Mackenzie, 1st Earl of Cromartie Yes
Archibald Primrose, 1st Earl of Rosebery Yes
David Boyle, 1st Earl of Glasgow Yes
Charles Hope, 1st Earl of Hopetoun LIKELY Linlithgowshire Yes
Henry Scott, 1st Earl of Deloraine Yes
Archibald Campbell, Earl of Illay Yes
William Hay, Viscount Dupplin Yes
William Forbes, 12th Lord Forbes Yes
John Elphinstone, 8th Lord Elphinstone Yes
William Ross, 12th Lord Ross Yes
James Sandilands, 7th Lord Torphichen Yes
Lord Fraser Yes
George Ogilvy, 3rd Lord Banff Yes
Alexander Murray, 4th Lord Elibank Yes
Kenneth Sutherland, 3rd Lord Duffus Yes
Robert Rollo, 4th Lord Rollo Stirlingshire Yes
James Murray, Lord Philiphaugh Lord Clerk Register/Selkirkshire Yes
Adam Cockburn, Lord Ormiston Lord Justice Clerk Yes
Sir Robert Dickson of Inverask Edinburghshire Yes
William Nisbet of Dirletoun Haddingtonshire Squadrone Volante Yes
John Cockburn, younger, of Ormestoun Haddingtonshire Squadrone Volante Yes
Sir John Swintoun of that ilk Berwickshire Court Party Yes
Sir Alexander Campbell of Cessnock Berwickshire Yes
Sir William Kerr of Greenhead Roxburghshire Squadrone Volante Yes
Archibald Douglas of Cavers Roxburghshire Court Party Yes
William Bennet of Grubbet Roxburghshire Court Party Yes
Mr John Murray of Bowhill Selkirkshire Court Party Yes
Mr John Pringle of Haining Selkirkshire Court Party Yes
William Morison of Prestongrange Peeblesshire Court Party Yes
Alexander Horseburgh of that ilk Peeblesshire Yes
George Baylie of Jerviswood Lanarkshire Squadrone Volante Yes
Sir John Johnstoun of Westerhall Dumfriesshire Court Party Yes
William Dowglass of Dornock Dumfriesshire Yes
Mr William Stewart of Castlestewart Wigtownshire Yes
Mr John Stewart of Sorbie Wigtownshire Court Party Yes
Mr Francis Montgomery of Giffan Ayrshire Court Party Yes
Mr William Dalrymple of Glenmuir Ayrshire Court Party Yes
Mr Robert Stewart of Tillicultrie Buteshire Yes
Sir Robert Pollock of that ilk Renfrewshire Court Party Yes
Mr John Montgomery of Wrae Linlithgowshire Yes
John Halden of Glenagies Perthshire Squadrone Volante Yes
Mongo Graham of Gorthie Perthshire Squadrone Volante Yes
Sir Thomas Burnet of Leyes Kincardineshire Court Party Yes
William Seton, younger, of Pitmedden Aberdeenshire Squadrone Volante Yes
Alexander Grant, younger, of that ilk Inverness-shire Court Party Yes
Sir William Mackenzie Yes
Mr Aeneas McLeod of Cadboll Cromartyshire Yes
Mr John Campbell of Mammore Argyllshire Court Party Yes
Sir James Campbell of Auchinbreck Argyllshire Court Party Yes
James Campbell, younger, of Ardkinglass Argyllshire Court Party Yes
Sir William Anstruther of that ilk Fife Yes
James Halyburton of Pitcurr Forfarshire Squadrone Volante Yes
Alexander Abercrombie of Glassoch Banffshire Court Party Yes
Mr James Dunbarr, younger, of Hemprigs Caithness Yes
Alexander Douglas of Eagleshay Orkney and Shetland Court Party Yes
Sir John Bruce, 2nd Baronet Kinross-shire Squadrone Volante Yes
John Scrimsour Dundee Yes
Lieutenant Colonel John Areskine Yes
John Mure Likely Ayr Yes
James Scott Montrose Court Party Yes
Sir John Anstruther, 1st Baronet, of Anstruther Anstruther Easter Yes
James Spittle Inverkeithing Yes
Mr Patrick Moncrieff Kinghorn Court Party Yes
Sir Andrew Home Kirkcudbright Squadrone Volante Yes
Sir Peter Halket Dunfermline Squadrone Volante Yes
Sir James Smollet Dumbarton Court Party Yes
Mr William Carmichell Lanark Yes
Mr William Sutherland Elgin Yes
Captain Daniel McLeod Tain Yes
Sir David Dalrymple, 1st Baronet Culross Court Party Yes
Sir Alexander Ogilvie Banff Yes
Mr John Clerk Whithorn Court Party Yes
John Ross Yes
Hew Dalrymple, Lord North Berwick North Berwick Yes
Mr Patrick Ogilvie Cullen Court Party Yes
George Allardyce Kintore Court Party Yes
William Avis Yes
Mr James Bethun Kilrenny Yes
Mr Roderick McKenzie Fortrose Yes
John Urquhart Dornoch Yes
Daniel Campbell Inveraray Court Party Yes
Sir Robert Forbes Inverurie Yes
Mr Robert Dowglass Kirkwall Yes
Mr Alexander Maitland Inverbervie Court Party Yes
Mr George Dalrymple Stranraer Yes
Mr Charles Campbell Campbeltown Yes
Total Ayes 106
James Hamilton, 4th Duke of Hamilton No
William Johnstone, 1st Marquess of Annandale Annan No
Charles Hay, 13th Earl of Erroll No
William Keith, 9th Earl Marischal No
David Erskine, 9th Earl of Buchan No
Alexander Sinclair, 9th Earl of Caithness No
John Fleming, 6th Earl of Wigtown No
James Stewart, 5th Earl of Galloway No
David Murray, 5th Viscount of Stormont No
William Livingston, 3rd Viscount of Kilsyth No
William Fraser, 12th Lord Saltoun No
Francis Sempill, 10th Lord Sempill No
Charles Oliphant, 7th Lord Oliphant No
John Elphinstone, 4th Lord Balmerino No
Walter Stuart, 6th Lord Blantyre Linlithgow No
William Hamilton, 3rd Lord Bargany Queensferry No
John Hamilton, 2nd Lord Belhaven and Stenton No
Lord Colvill No
Patrick Kinnaird, 3rd Lord Kinnaird No
Sir John Lawder of Fountainhall Haddingtonshire No
Andrew Fletcher of Saltoun Haddingtonshire No
Sir Robert Sinclair, 3rd Baronet Berwickshire No
Sir Patrick Home of Rentoun Berwickshire No
Sir Gilbert Elliot of Minto Roxburghshire No
William Bayllie of Lamingtoun Lanarkshire No
John Sinclair, younger, of Stevensone Lanarkshire No
James Hamilton of Aikenhead Lanarkshire No
Mr Alexander Fergusson of Isle Dumfriesshire No
Sir Hugh Cathcart of Carletoun Ayrshire No
John Brisbane, younger, of Bishoptoun Ayrshire No
Mr William Cochrane of Kilmaronock Dumbartonshire No
Sir Humphray Colquhoun of Luss Dumbartonshire No
Sir John Houstoun of that ilk Renfrewshire No
Robert Rollo of Powhouse No
Thomas Sharp of Houstoun Linlithgowshire No
John Murray of Strowan No
Alexander Gordon of Pitlurg Aberdeenshire No
John Forbes of Colloden Nairnshire No
David Bethun of Balfour Fife No
Major Henry Balfour of Dunboog Fife No
Mr Thomas Hope of Rankeillor No
Mr Patrick Lyon of Auchterhouse Forfarshire No
Mr James Carnagie of Phinhaven Forfarshire No
David Graham, younger, of Fintrie Forfarshire No
William Maxwell of Cardines Kirkcudbrightshire No
Alexander McKye of Palgown Kirkcudbrightshire No
James Sinclair of Stempster Caithness No
Sir Henry Innes, younger, of that ilk Elginshire No
Mr George McKenzie of Inchcoulter Ross-shire No
Robert Inglis Edinburgh No
Alexander Robertson Perth No
Walter Stewart No
Hugh Montgomery Glasgow Court Party No
Alexander Edgar Haddington No
Alexander Duff Banffshire No
Francis Molison Brechin No
Walter Scott Jedburgh No
Robert Scott Selkirk No
Robert Kellie Dunbar No
John Hutchesone Arbroath No
Archibald Scheills Peebles No
Mr John Lyon Forfar No
George Brodie Forres No
George Spens Rutherglen No
Sir David Cuningham Lauder No
Mr John Carruthers Lochmaben No
George Home New Galloway No
John Bayne Dingwall No
Mr Robert Fraser Wick No
Total Noes 69
Total Votes 175
Sources: Records of the Parliament of Scotland, Parliamentary Register, p.598
See also[edit]
Acts of Union 1800
History of democracy
List of treaties
MacCormick v Lord Advocate
Parliament of the United Kingdom
Political union
Real union
English independence
Scottish independence
Unionism in Scotland
Welsh independence
Notes[edit]
Jump up ^ The citation of this Act by this short title was authorised by section 1 of, and Schedule 1 to, the Short Titles Act 1896. Due to the repeal of those provisions, it is now authorised by section 19(2) of the Interpretation Act 1978.
Jump up ^ Article I of the Treaty of Union
Jump up ^ Act of Union 1707, Article 3
Jump up ^ Simon Schama (presenter) (22 May 2001). "Britannia Incorporated". A History of Britain. Episode 10. 3 minutes in. BBC One.
Jump up ^ ABDN.ac.uk
Jump up ^ Larkin, James F.; Hughes, Paul L., eds. (1973). Stuart Royal Proclamations: Volume I. Clarendon Press. p. 19.
Jump up ^ Lockyer, R. (1998). James VI and I. London: Addison Wesley Longman. pp. 51–52. ISBN 0-582-27962-3.
Jump up ^ Lockyer, op. cit., pp. 54–59
Jump up ^ Parliament.uk Archived 10 December 2008 at the Wayback Machine.
Jump up ^ Lockyer, op. cit., p.59
^ Jump up to: a b Parliament.uk Archived 12 October 2008 at the Wayback Machine.
Jump up ^ Constitution.org
Jump up ^ The 1657 Act's long title was An Act and Declaration touching several Acts and Ordinances made since 20 April 1653, and before 3 September 1654, and other Acts
Jump up ^ C. Whatley, op. cit., p.95
Jump up ^ C. Whatley, op. cit., p.30
Jump up ^ Whatley, C. (2006). The Scots and the Union. Edinburgh: Edinburgh University Press. p. 91. ISBN 0-7486-1685-3.
^ Jump up to: a b "The commissioners". UK Parliament website. 2007. Archived from the original on 19 June 2009. Retrieved 5 February 2013.
Jump up ^ "The course of negotiations". UK Parliament website. 2007. Archived from the original on 21 July 2009. Retrieved 5 February 2013.
Jump up ^ "Ratification". UK parliament website. 2007. Archived from the original on 19 June 2009. Retrieved 5 February 2013.
Jump up ^ "1 May 1707 – the Union comes into effect". UK Parliament website. 2007. Archived from the original on 19 June 2009. Retrieved 5 February 2013.
Jump up ^ Whatley, C. A. (2001). Bought and sold for English Gold? Explaining the Union of 1707. East Linton: Tuckwell Press. p. 48. ISBN 1-86232-140-X.
Jump up ^ Watt, Douglas. The Price of Scotland: Darien, Union and the wealth of nations. Luath Press 2007.
Jump up ^ Parliament.uk Archived 25 September 2008 at the Wayback Machine.
Jump up ^ Bambery, Chris (2014). A People's History of Scotland. Verso.
^ Jump up to: a b "Scottish Referendums". Bbc.co.uk. Retrieved 2016-03-16.
Jump up ^ The Humble Address of the Commissioners to the General Convention of the Royal Burrows of this Ancient Kingdom Convened the Twenty-Ninth of October 1706, at Edinburgh
Jump up ^ Notes by John Purser to CD Scotland's Music, Facts about Edinburgh.
Jump up ^ Riley, P. J. W. (1969). "The Union of 1707 as an Episode in English Politics". The English Historical Review. 84 (332): 498–527 [pp. 523–524]. JSTOR 562482. doi:10.1093/ehr/lxxxiv.cccxxxii.498.
Jump up ^ G.N. Clark, The Later Stuarts, 1660-1714 (2nd ed. 1956) pp 290-93.
Jump up ^ Gordon Brown (2014). My Scotland, Our Britain: A Future Worth Sharing. Simon & Schuster UK. p. 150.
Jump up ^ House of Lords – Written answers, 6 November 2006, TheyWorkForYou.com
Jump up ^ Announced by the Scottish Culture Minister, Patricia Ferguson, 9 November 2006
Further reading[edit]
Bowie, Karin. Scottish Public Opinion and the Anglo-Scottish Union, 1699-1707 (Boydell Press, 2007).
Brown, Stewart J. and Christopher A. Whatley, eds. The Union of 1707: New Dimensions (Edinburgh UP, 2008).
Deschamps, Yannick. "Résistances écossaises à l’union de 1707: essai historiographique," Dix-huitième siècle (2012) #44 pp 601-20 DOI : 10.3917/dhs.044.0601; online--use built in translator in Chrome for English
Ferguson, William. "The Making of the Treaty of Union of 1707" Scottish Historical Review 43, (1964), p. 89-110.
Ferguson, William. Scotland’s Relations with England: A Survey to 1707 (Edinburgh John Donald, 1977), pp. 180-277.
Herman, Arthur. How the Scots Invented the Modern World. Three Rivers Press, 2001. ISBN 0-609-80999-7
Riley, P. W. J. The Union of England and Scotland: A Study in Anglo-Scottish Politics of the Eighteenth Century (Manchester UP, 1978).
Smout, T. C. "Anglo-Scottish Union of 1707. I. The Economic Background" Economic History Review (1964) 16:3, pp. 455-467.
Stephen, Jeffrey. Scottish Presbyterians and the Act of Union 1707 (Edinburgh UP, 2007).
Other books[edit]
Defoe, Daniel. A tour thro' the Whole Island of Great Britain, 1724–27
Defoe, Daniel. The Letters of Daniel Defoe, GH Healey editor. Oxford: 1955.
Fletcher, Andrew (Saltoun). An Account of a Conversation
Lockhart, George, "The Lockhart Papers", 1702–1728
External links[edit]
Wikisource has original text related to this article:
Acts of Union 1707
Union with England Act and Union with Scotland Act – Full original text
Treaty of Union and the Darien Experiment, University of Guelph, McLaughlin Library, Library and Archives Canada
Text of the Union with Scotland Act 1706 as in force today (including any amendments) within the United Kingdom, from legislation.gov.uk
Text of the Union with England Act 1707 as in force today (including any amendments) within the United Kingdom, from legislation.gov.uk
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Chromosomal crossover
From Wikipedia, the free encyclopedia
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Crossing over occurs between prophase 1 and metaphase 1 and is the process where homologous chromosomes pair up with each other and exchange different segments of their genetic material to form recombinant chromosomes. It can also happen during mitotic division,[1] which may result in loss of heterozygosity. Crossing over is essential for the normal segregation of chromosomes during meiosis.[citation needed] Crossing over also accounts for genetic variation, because due to the swapping of genetic material during crossing over, the chromatids held together by the centromere are no longer identical. So, when the chromosomes go on to meiosis II and separate, some of the daughter cells receive daughter chromosomes with recombined alleles. Due to this genetic recombination, the offspring have a different set of alleles and genes than their parents do. In the diagram, genes B and b are crossed over with each other, making the resulting recombinants after meiosis Ab, AB, ab, and aB.
Thomas Hunt Morgan's illustration of crossing over (1916)
A double crossing over
Chromosomal crossover (or crossing over) is the exchange of genetic material between homologous chromosomes that results in recombinant chromosomes during sexual reproduction. It is one of the final phases of genetic recombination, which occurs in the pachytene stage of prophase I of meiosis during a process called synapsis. Synapsis begins before the synaptonemal complex develops and is not completed until near the end of prophase I. Crossover usually occurs when matching regions on matching chromosomes break and then reconnect to the other chromosome.
Crossing over was described, in theory, by Thomas Hunt Morgan. He relied on the discovery of Frans Alfons Janssens who described the phenomenon in 1909 and had called it "chiasmatypie". The term chiasma is linked, if not identical, to chromosomal crossover. Morgan immediately saw the great importance of Janssens' cytological interpretation of chiasmata to the experimental results of his research on the heredity of Drosophila. The physical basis of crossing over was first demonstrated by Harriet Creighton and Barbara McClintock in 1931.[2]
The linked frequency of crossing over between two gene loci (markers) is the crossing-over value . For fixed set of genetic and environmental conditions, recombination in a particular region of a linkage structure (chromosome) tends to be constant and the same is then true for the crossing-over value which is used in the production of genetic maps.[3][4]
Contents
[hide]
1 Origins
1.1 DNA repair theory
1.2 Links to bacterial transformation
2 Chemistry
2.1 MSH4 and MSH5
2.2 Chiasma
3 Consequences
4 Non-homologous crossover
5 See also
6 References
Origins[edit]
There are two popular and overlapping theories that explain the origins of crossing-over, coming from the different theories on the origin of meiosis. The first theory rests upon the idea that meiosis evolved as another method of DNA repair, and thus crossing-over is a novel way to replace possibly damaged sections of DNA.[5] The second theory comes from the idea that meiosis evolved from bacterial transformation, with the function of propagating diversity.[6]
DNA repair theory[edit]
Crossing over and DNA repair are very similar processes, which utilize many of the same protein complexes.[5][7][8] Recombinases and primases lay a foundation of nucleotides along the DNA sequence. One such particular protein complex that is conserved between processes is RAD51, a well conserved recombinase protein that has been shown to be crucial in DNA repair as well as cross over.[9] Several other genes in D. melanogaster have been linked as well to both processes, by showing that mutants at these specific loci cannot undergo DNA repair or crossing over. Such genes include mei-41, mei-9, hdm, spnA, and brca2.[5] This large group of conserved genes between processes supports the theory of a close evolutionary relationship. Furthermore, DNA repair and crossover have been found to favor similar regions on chromosomes. In an experiment using radiation hybrid mapping on wheat’s (Triticum aestivum L.) 3B chromosome, crossing over and DNA repair were found to occur predominantly in the same regions.[10] Furthermore, crossing over has been correlated to occur in response to stressful, and likely DNA damaging, conditions [11][12]
Links to bacterial transformation[edit]
The process of bacterial transformation also shares many similarities with chromosomal cross over, particularly in the formation of overhangs on the sides of the broken DNA strand, allowing for the annealing of a new strand. Bacterial transformation itself has been linked to DNA repair many times.[5] The second theory comes from the idea that meiosis evolved from bacterial transformation, with the function of propagating genetic diversity.[6] .[13] Thus, this evidence suggests that it is a question of whether cross over is linked to DNA repair or bacterial transformation, as the two do not appear to be mutually exclusive. It is likely that crossing over may have evolved from bacterial transformation, which in turn developed from DNA repair, thus explaining the links between all three processes.
Chemistry[edit]
A current model of meiotic recombination, initiated by a double-strand break or gap, followed by pairing with an homologous chromosome and strand invasion to initiate the recombinational repair process. Repair of the gap can lead to crossover (CO) or non-crossover (NCO) of the flanking regions. CO recombination is thought to occur by the Double Holliday Junction (DHJ) model, illustrated on the right, above. NCO recombinants are thought to occur primarily by the Synthesis Dependent Strand Annealing (SDSA) model, illustrated on the left, above. Most recombination events appear to be the SDSA type.
Meiotic recombination may be initiated by double-stranded breaks that are introduced into the DNA by exposure to DNA damaging agents[5] or the Spo11 protein.[14] One or more exonucleases then digest the 5’ ends generated by the double-stranded breaks to produce 3’ single-stranded DNA tails (see diagram). The meiosis-specific recombinase Dmc1 and the general recombinase Rad51 coat the single-stranded DNA to form nucleoprotein filaments.[15] The recombinases catalyze invasion of the opposite chromatid by the single-stranded DNA from one end of the break. Next, the 3’ end of the invading DNA primes DNA synthesis, causing displacement of the complementary strand, which subsequently anneals to the single-stranded DNA generated from the other end of the initial double-stranded break. The structure that results is a cross-strand exchange, also known as a Holliday junction. The contact between two chromatids that will soon undergo crossing-over is known as a chiasma. The Holliday junction is a tetrahedral structure which can be 'pulled' by other recombinases, moving it along the four-stranded structure.
Holliday Junction
Molecular structure of a Holliday junction.
Molecular structure of a Holliday junction. From PDB: 3CRX.
MSH4 and MSH5[edit]
The MSH4 and MSH5 proteins form a hetero-oligomeric structure (heterodimer) in yeast and humans.[16][17][18] In the yeast Saccharomyces cerevisiae MSH4 and MSH5 act specifically to facilitate crossovers between homologous chromosomes during meiosis.[16] The MSH4/MSH5 complex binds and stabilizes double Holliday junctions and promotes their resolution into crossover products. An MSH4 hypomorphic (partially functional) mutant of S. cerevisiae showed a 30% genome wide reduction in crossover numbers, and a large number of meioses with non exchange chromosomes.[19] Nevertheless, this mutant gave rise to spore viability patterns suggesting that segregation of non-exchange chromosomes occurred efficiently. Thus in S. cerevisiae proper segregation apparently does not entirely depend on crossovers between homologous pairs.
Chiasma[edit]
The grasshopper Melanoplus femur-rubrum was exposed to an acute dose of X-rays during each individual stage of meiosis, and chiasma frequency was measured.[20] Irradiation during the leptotene-zygotene stages of meiosis (that is, prior to the pachytene period in which crossover recombination occurs) was found to increase subsequent chiasma frequency. Similarly, in the grasshopper Chorthippus brunneus, exposure to X-irradiation during the zygotene-early pachytene stages caused a significant increase in mean cell chiasma frequency.[21] Chiasma frequency was scored at the later diplotene-diakinesis stages of meiosis. These results suggest that X-rays induce DNA damages that are repaired by a crossover pathway leading to chiasma formation.
Consequences[edit]
The difference between gene conversion and chromosomal crossover.
In most eukaryotes, a cell carries two versions of each gene, each referred to as an allele. Each parent passes on one allele to each offspring. An individual gamete inherits a complete haploid complement of alleles on chromosomes that are independently selected from each pair of chromatids lined up on the metaphase plate. Without recombination, all alleles for those genes linked together on the same chromosome would be inherited together. Meiotic recombination allows a more independent segregation between the two alleles that occupy the positions of single genes, as recombination shuffles the allele content between homologous chromosomes.
Recombination results in a new arrangement of maternal and paternal alleles on the same chromosome. Although the same genes appear in the same order, some alleles are different. In this way, it is theoretically possible to have any combination of parental alleles in an offspring, and the fact that two alleles appear together in one offspring does not have any influence on the statistical probability that another offspring will have the same combination. This principle of "independent assortment" of genes is fundamental to genetic inheritance.[22] However, the frequency of recombination is actually not the same for all gene combinations. This leads to the notion of "genetic distance", which is a measure of recombination frequency averaged over a (suitably large) sample of pedigrees. Loosely speaking, one may say that this is because recombination is greatly influenced by the proximity of one gene to another. If two genes are located close together on a chromosome, the likelihood that a recombination event will separate these two genes is less than if they were farther apart. Genetic linkage describes the tendency of genes to be inherited together as a result of their location on the same chromosome. Linkage disequilibrium describes a situation in which some combinations of genes or genetic markers occur more or less frequently in a population than would be expected from their distances apart. This concept is applied when searching for a gene that may cause a particular disease. This is done by comparing the occurrence of a specific DNA sequence with the appearance of a disease. When a high correlation between the two is found, it is likely that the appropriate gene sequence is really closer.[23]
Non-homologous crossover[edit]
Crossovers typically occur between homologous regions of matching chromosomes, but similarities in sequence and other factors can result in mismatched alignments. Most DNA is composed of base pair sequences repeated very large numbers of times.[24] These repetitious segments, often referred to as satellites, are fairly homogenous among a species.[24] During DNA replication, each strand of DNA is used as a template for the creation of new strands using a partially-conserved mechanism; proper functioning of this process results in two identical, paired chromosomes, often called sisters. Sister chromatid crossover events are known to occur at a rate of several crossover events per cell per division in eukaryotes. [24] Most of these events involve an exchange of equal amounts of genetic information, but unequal exchanges may occur due to sequence mismatch. These are referred to by a variety of names, including non-homologous crossover, unequal crossover, and unbalanced recombination, and result in an insertion or deletion of genetic information into the chromosome. While rare compared to homologous crossover events, these mutations are drastic, affecting many loci at the same time. They are considered the main driver behind the generation of gene duplications and are a general source of mutation within the genome.[25]
Many proteins are implicated in DSB repair; this figure shows a probable, simplified mechanism.
The specific causes of non-homologous crossover events are unknown, but several influential factors are known to increase the likelihood of an unequal crossover. One common vector leading to unbalanced recombination is the repair of double-strand breaks (DSBs).[26] DSBs are often repaired using non-homologous end joining, a process which involves invasion of a template strand by the DSB strand (see figure below). Nearby homologous regions of the template strand are often used for repair, which can give rise to either insertions or deletions in the genome if a non-homologous but complementary part of the template strand is used.[26] Sequence similarity is a major player in crossover – crossover events are more likely to occur in long regions of close identity on a gene.[27] This means that any section of the genome with long sections of repetitive DNA is prone to crossover events.
The presence of transposable elements is another influential element of non-homologous crossover. Repetitive regions of code characterize transposable elements; complementary but non-homologous regions are ubiquitous within transposons. Because chromosomal regions composed of transposons have large quantities of identical, repetitious code in a condensed space, it is thought that transposon regions undergoing a crossover event are more prone to erroneous complementary match-up;[28] that is to say, a section of a chromosome containing a lot of identical sequences, should it undergo a crossover event, is less certain to match up with a perfectly homologous section of complementary code and more prone to binding with a section of code on a slightly different part of the chromosome. This results in unbalanced recombination, as genetic information may be either inserted or deleted into the new chromosome, depending on where the recombination occurred.
While the motivating factors behind unequal recombination remain obscure, elements of the physical mechanism have been elucidated. Mismatch repair (MMR) proteins, for instance, are a well-known regulatory family of proteins, responsible for regulating mismatched sequences of DNA during replication and escape regulation.[29] The operative goal of MMRs is the restoration of the parental genotype. One class of MMR in particular, MutSβ, is known to initiate the correction of insertion-deletion mismatches of up to 16 nucleotides.[29] Little is known about the excision process in eukaryotes, but E. coli excisions involve the cleaving of a nick on either the 5’ or 3’ strand, after which DNA helicase and DNA polymerase III bind and generate single-stranded proteins, which are digested by exonucleases and attached to the strand by ligase.[29] Multiple MMR pathways have been implicated in the maintenance of complex organism genome stability, and any of many possible malfunctions in the MMR pathway result in DNA editing and correction errors.[30] Therefore, while it is not certain precisely what mechanisms lead to errors of non-homologous crossover, it is extremely likely that the MMR pathway is involved.
See also[edit]
Unequal crossing over
Coefficient of coincidence
Genetic distance
Independent assortment
Mitotic crossover
Recombinant frequency
References[edit]
Jump up ^ Griffiths, AJF; Gelbart, WM; Miller, JH; et al. (1999). "Modern Genetic Analysis: Mitotic Crossing-Over". New York: W. H. Freeman.
Jump up ^ Creighton H, McClintock B (1931). "A Correlation of Cytological and Genetical Crossing-Over in Zea Mays". Proc Natl Acad Sci USA. 17 (8): 492–7. doi:10.1073/pnas.17.8.492. PMC 1076098  . PMID 16587654. (Original paper)
Jump up ^ Rieger R. Michaelis A., Green M. M. (1976). Glossary of genetics and cytogenetics: Classical and molecular. Heidelberg - New York: Springer-Verlag. ISBN 3-540-07668-9. CS1 maint: Uses authors parameter (link)
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^ Jump up to: a b c d e Harris Bernstein, Carol Bernstein and Richard E. Michod (2011). Meiosis as an Evolutionary Adaptation for DNA Repair. Chapter 19 in DNA Repair. Inna Kruman, editor. InTech Open Publisher. doi:10.5772/25117 http://www.intechopen.com/books/dna-repair/meiosis-as-an-evolutionary-adaptation-for-dna-repair
^ Jump up to: a b Bernstein, H; Bernstein, C (2010). "Evolutionary origin of recombination during meiosis". BioScience. 60 (7): 498–505. doi:10.1525/bio.2010.60.7.5.
Jump up ^ Dangel, NJ; Knoll, A; Puchta, H (2014). "MHF1 plays Fanconi anaemia complementation group M protein (FANCM)-dependent and FANCM-independent roles in DNA repair and homologous recombination in plants". Plant J. 78 (5): 822–33. doi:10.1111/tpj.12507. PMID 24635147.
Jump up ^ Saponaro, M; Callahan, D; Zheng, X; Liberi, G (2010). "Cdk1 Targets Srs2 to Complete Synthesis-Dependent Strand Annealing and to Promote Recombinational Repair". PLoS Genet. 6 (2): e1000858. doi:10.1371/journal.pgen.1000858. Retrieved 20 March 2015.
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Jump up ^ Charpentier, X (2010). "Antibiotics and UV Radiation Induce Competence for Natural Transformation in Legionella pneumophila". Journal of Bacteriology. 193 (5): 1114–1121. doi:10.1128/JB.01146-10. PMC 3067580  . PMID 21169481. Retrieved 14 March 2015.
Jump up ^ Keeney, S; Giroux, CN; Kleckner, N (1997). "Meiosis-Specific DNA Double-Strand Breaks Are Catalyzed by Spo11, a Member of a Widely Conserved Protein Family". Cell. 88 (3): 375–84. doi:10.1016/S0092-8674(00)81876-0. PMID 9039264.
Jump up ^ Sauvageau, S; Stasiak, Az; Banville, I; Ploquin, M; Stasiak, A; Masson, Jy (June 2005). "Fission Yeast Rad51 and Dmc1, Two Efficient DNA Recombinases Forming Helical Nucleoprotein Filaments" (Free full text). Molecular and Cellular Biology. 25 (11): 4377–87. doi:10.1128/MCB.25.11.4377-4387.2005. ISSN 0270-7306. PMC 1140613  . PMID 15899844.
^ Jump up to: a b Pochart P, Woltering D, Hollingsworth NM (1997). "Conserved properties between functionally distinct MutS homologs in yeast". J. Biol. Chem. 272 (48): 30345–9. doi:10.1074/jbc.272.48.30345. PMID 9374523.
Jump up ^ Winand NJ, Panzer JA, Kolodner RD (1998). "Cloning and characterization of the human and Caenorhabditis elegans homologs of the Saccharomyces cerevisiae MSH5 gene". Genomics. 53 (1): 69–80. doi:10.1006/geno.1998.5447. PMID 9787078.
Jump up ^ Bocker T, Barusevicius A, Snowden T, Rasio D, Guerrette S, Robbins D, Schmidt C, Burczak J, Croce CM, Copeland T, Kovatich AJ, Fishel R (1999). "hMSH5: a human MutS homologue that forms a novel heterodimer with hMSH4 and is expressed during spermatogenesis". Cancer Res. 59 (4): 816–22. PMID 10029069.
Jump up ^ Krishnaprasad GN, Anand MT, Lin G, Tekkedil MM, Steinmetz LM, Nishant KT (2015). "Variation in crossover frequencies perturb crossover assurance without affecting meiotic chromosome segregation in Saccharomyces cerevisiae". Genetics. 199 (2): 399–412. doi:10.1534/genetics.114.172320. PMC 4317650  . PMID 25467183.
Jump up ^ Church K, Wimber DE (1969). "Meiosis in the grasshopper: chiasma frequency after elevated temperature and x-rays". Can. J. Genet. Cytol. 11 (1): 209–16. doi:10.1139/g69-025. PMID 5797806.
Jump up ^ Westerman M (1971). "The effect of x-irradiation on chiasma frequency in Chorthippus brunneus". Heredity. 27 (1): 83–91. doi:10.1038/hdy.1971.73. PMID 5289295.
Jump up ^ "genetic recombination".
Jump up ^ Genetic Recombination
^ Jump up to: a b c Smith, George P. (1976-01-01). "Evolution of Repeated DNA Sequences by Unequal Crossover". Science. 191 (4227): 528–535. doi:10.1126/science.1251186. JSTOR 1741301.
Jump up ^ Graur, Dan; Li, Wen-Hsiung (2000-01-01). Fundamentals of Molecular Evolution. Sinauer. ISBN 9780878932665.
^ Jump up to: a b Puchta, Holger (2005-01-01). "The repair of double-strand breaks in plants: mechanisms and consequences for genome evolution". Journal of Experimental Botany. 56 (409): 1–14. doi:10.1093/jxb/eri025. ISSN 0022-0957. PMID 15557293.
Jump up ^ Metzenberg, Ab; et al. (May 1991). "Homology Requirements for Unequal Crossing Over in Humans". Genetics.
Jump up ^ Robberecht, Caroline; Voet, Thierry; Esteki, Masoud Zamani; Nowakowska, Beata A.; Vermeesch, Joris R. (2012-12-03). "Non-allelic homologous recombination between retrotransposable elements is a driver of de novo unbalanced translocations". Genome Research. 23: gr.145631.112. doi:10.1101/gr.145631.112. ISSN 1088-9051. PMC 3589530  . PMID 23212949.
^ Jump up to: a b c Kunkel, Thomas A.; Erie, Dorothy A. (2005-01-01). "Dna Mismatch Repair". Annual Review of Biochemistry. 74 (1): 681–710. doi:10.1146/annurev.biochem.74.082803.133243. PMID 15952900.
Jump up ^ Surtees, J. A., J.A.; Argueso, J.L.; Alani, E. (2004). "Mismatch Repair Proteins: Key Regulators of Genetic Recombination". Cytogenetic and Genome Research. 107: 146–59. doi:10.1159/000080593.
[hide]
v
t
e
Genetics: homologous recombination / mobile genetic elements
Primarily prokaryotic
Conjugation
Transduction
Transformation
Occurs in eukaryotes
Transfection
Chromosomal crossover
Gene conversion
Fusion gene
Horizontal gene transfer
Sister chromatid exchange
Transposon
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