Field Marshal William Joseph Slim, 1st Viscount Slim,  (6 August 1891 – 14 December 1970), usually known as Bill Slim, was a British military commander and the 13th Governor-General of Australia.

Slim saw active service in both the First and Second World Wars and was wounded in action three times. During the Second World War he led the Fourteenth Army, the so-called "forgotten army" in the Burma campaign. After the war he became the first British officer who had served in the Indian Army to be appointed Chief of the Imperial General Staff. From 1953 to 1959 he was Governor-General of Australia.

In the early 1930s, Slim also wrote novels, short stories, and other publications under the pen name Anthony Mills.

Early years
William Slim was born at 72 Belmont Road, St Andrews, Bristol, the son of John Slim by his marriage to Charlotte Tucker, and was baptised there at St Bonaventure's Roman Catholic church, Bishopston. He was brought up first in Bristol, attending St Bonaventure's Primary School, then St Brendan's College, before moving to Birmingham in his teens.  In Birmingham, he attended St Philip's Grammar School, Edgbaston and King Edward's School, Birmingham.

After leaving school, his father's failure in business as a wholesale ironmonger meant that the family could afford to send only one son, Slim's older brother, to the University of Birmingham, so between 1910 and 1914 Slim taught in a primary school and worked as a clerk in Stewarts & Lloyds, a metal-tube maker.

First World War
Despite having no other connection to the university, in 1912 Slim joined the Birmingham University Officers' Training Corps, and he was thus able to be commissioned as a temporary second lieutenant into the Royal Warwickshire Regiment on 22 August 1914, on the outbreak of the First World War; in later life, as a result of his modest social origins and his unpretentious manner, he was sometimes wrongly supposed to have risen from the ranks. He was badly wounded at Gallipoli. On return to England, he was granted a regular commission as a second lieutenant in the West India Regiment.

In October 1916, he rejoined the Royal Warwickshire Regiment in Mesopotamia. On 4 March 1917, he was promoted to lieutenant (with seniority back-dated to October 1915). He was wounded a second time in 1917. Having been previously given the temporary rank of captain, he was awarded the Military Cross on 7 February 1918 for actions in Mesopotamia.

Evacuated to India, he was given the temporary rank of major in the 6th Gurkha Rifles on 2 November 1918. He was formally promoted to captain and transferred to the Indian Army on 22 May 1919.

Interwar career
Slim became battalion adjutant with the 6th Gurkha Rifles in 1921.

On 1 January 1926, he married Aileen Robertson, daughter of Rev John Anderson Robertson (d.1941) minister of Cramond near Edinburgh. They had one son and one daughter. Later that year Slim was sent to the Staff College, Quetta. On 5 June 1929, he was appointed a General Staff Officer, Second Grade.

On 1 January 1930, he was given the brevet rank of major, with formal promotion to this rank made on 19 May 1933. His performance at Staff College resulted in his appointment first to Army Headquarters India in Delhi and then to Staff College, Camberley, in England (as a General Staff Officer, Second Grade), where he taught from 1934 to 1937. During this period, he also wrote novels, short stories, and other publications under the pen name of Anthony Mills, in order to further his literary interests, as well as to supplement his then modest army salary.

He attended the Imperial Defence College in 1937. The following year he was promoted to lieutenant colonel and given command of the 2nd Battalion, 7th Gurkha Rifles. In 1939 he was briefly given the temporary rank of brigadier as commander of his battalion. On 8 June 1939, he was promoted to colonel (again with temporary rank of brigadier) and appointed head of the Senior Officers' School, Belgaum in India.

Second World War

East African Campaign
On the outbreak of the Second World War, Slim was given command of the 10th Indian Infantry Brigade of the 5th Indian Infantry Division and was sent to Sudan. He took part in the East African campaign to liberate Ethiopia from the Italians. Slim was wounded again in Eritrea. On 21 January 1941, he was hit when his vehicle was strafed by Fiat CR.42 fighters during the advance on Agordat.

Middle East
Recovering from his wounds but still unfit for active service, Slim was temporarily employed on the General Staff at GHQ in Delhi. He was involved in the planning for potential operations in Iraq where trouble was expected. By early May 1941 Slim had been appointed Brigadier General Staff (chief staff officer) to Edward Quinan the commander designate for operations in Iraq, arriving in Basra on 7 May. Not long afterwards, Major-General Fraser, commanding the 10th Indian Infantry Division, fell ill and was relieved of his command, and Slim was promoted to take his place on 15 May 1941 with the acting rank of major-general.

He led the Indian 10th Infantry Division as part of Iraqforce during the Anglo-Iraqi War, the Syria–Lebanon campaign (where the division advanced up the river Euphrates to capture Deir ez-Zor), and the invasion of Persia. He was twice mentioned in despatches during 1941.

Burma campaign

In March 1942, Slim was given command of Burma Corps, also known as BurCorps, consisting of the 17th Indian Infantry Division and 1st Burma Division. Slim was made acting lieutenant general on 8 May 1942. The corps was under attack in Burma by the Japanese and, heavily outclassed by the more mobile and flexible Japanese, was soon forced to withdraw to India. On 28 October 1942, Slim was appointed a Commander of the Order of the British Empire (CBE).

Slim then took over XV Corps under the command of the Eastern Army. His command covered the coastal approaches from Burma to India, east of Chittagong. He had a series of disputes with Noel Irwin, commander of Eastern Army and, as a result, Irwin (although an army commander) took personal control of the initial advance by XV Corps into the Arakan Peninsula. The operations ended in disaster, during which Slim was restored to command of XV Corps, albeit too late to salvage the situation. Generals Irwin and Slim blamed each other for the result, but in the end Irwin was removed from his command, and Slim was promoted to command the new Fourteenth Army—formed from IV Corps (United Kingdom) (Imphal), XV Corps (Arakan) and XXXIII Corps (reserve) – later joined by XXXIV Corps. On 14 January 1943, Slim was awarded the Distinguished Service Order (DSO) for his actions in the Middle East during 1941. The American historians Alan Millet and Williamson Murray described Slim as:

Slim quickly got on with the task of training his new army to take the fight to the enemy. His basic premise was that off-road mobility was paramount: much heavy equipment was exchanged for mule- or air-transported equipment, and motor transport was kept to a minimum and restricted to vehicles that could cope with some of the worst combat terrain on Earth. The new doctrine dictated that if the Japanese had cut the lines of communication, then they too were surrounded. All units were to form defensive 'boxes', to be resupplied by air and assisted by integrated close air support and armour. The boxes were designed as an effective response to the tactics of infiltration practised by the Japanese in the war. Slim also supported increased offensive patrolling and night training, to encourage his soldiers to lose both their fear of the jungle and their belief that Japanese soldiers were better jungle fighters. He also instructed them to hold their positions once outflanked.

The Chin Hills formed a natural defensive barrier into Burma, which Slim would have preferred to outflank by an amphibious operation by landing further down the coast of Burma, but demands of the war in Europe meant the necessary landing craft were not available, forcing Slim to devise plans for advancing into Burma overland through the Chin hills. At the same time, the Japanese 15th Army, which formed the main striking force of the Burma Area Army, had grown from four divisions at the beginning of 1943 to eight divisions by the end of 1943 as the Japanese made preparations for invading India, which increased the difficulties of an overland advance into Burma. By 1943, the Burma Railway, which cost the lives of thousands of prisoners of war who built it, was finished; this allowed the Japanese to reinforce the Burma Area Army, and made invading India possible.

As Slim went about training his men for the rigours of jungle warfare, he clashed with Brigadier Orde Wingate, who took away some of Slim's best Gurkha, British and African units for his Chindit raiding group. Slim argued against the loss of his better units to Wingate, and maintained that though Wingate had a successful career in Palestine and Ethiopia he would discover that the Japanese were a considerably tougher foe than the Palestinians and the Italians that Wingate had hitherto been fighting. However, Slim did approve of Wingate's plans for aid to the hill tribes of Burma. The various hill peoples of Burma such as the Kachins, Karens, Chin, Nagas and the Shan collectively amounted to about 7 million of Burma's 17 million people, and unlike the Bamars, who had welcomed the Japanese as liberators, had stayed loyal to the British when the Japanese invaded. The hill peoples of Burma had suffered under Japanese rule, and were more than willing to wage guerrilla warfare against them. Slim approved of the plans of the SOE and OSS to provide arms and training to the hill tribes as a way to tie down Japanese forces that would otherwise be deployed against him.

At the start of 1944, Slim held the substantive rank of colonel with a war substantive rank of major-general and the acting, then upgraded to temporary, rank of lieutenant-general. In January 1944, when the Second Arakan Offensive was met by a Japanese counter-offensive, the Indian 7th Infantry Division was quickly surrounded along with parts of the Indian 5th Infantry Division and the 81st (West Africa) Division. The 7th Indian Division's defence was based largely on the "Admin Box" formed initially from drivers, cooks and suppliers. They were supplied by air, thus negating the importance of their lost supply lines. The Japanese forces were able to halt the offensive into Arakan but were unable to decisively defeat the allied forces or advance beyond the surrounded formations.

In early 1944, the Japanese Prime Minister General Hideki Tojo approved of plans for victory in Asia, calling for two operations, Operation U-Go as the invasion of India was code-named and Operation Ichi-Go which was intended to defeat China once and for all. The two operations in India and China were closely linked given that American supplies for China were flown over "the Hump" of the Himalayas and the Japanese wanted to take the Indian province of Assam in part to close the American air bases in India that sustained China at the same time that they were launching Operation Ichi-Go, the biggest Japanese offensive of all time, involving 2 million men. The Japanese knew that they lacked the logistics to invade India, and the plans for U-Go were based on the assumption that the British Fourteenth Army would just collapse, allowing the Japanese 15th Army to capture enough food to prevent its men from starving to death. Following the Japanese 15th Army into India was the Indian National Army commanded by Subhas Chandra Bose, an ardent nationalist. The Japanese believed that the mere presence of Bose in India would inspire the men of the Indian Army to mutiny and murder their British officers, and set off an anti-British revolution that would allow the Japanese 15th Army to take all of India.

Slim was appointed a Companion of the Order of the Bath (CB) in the 1944 New Year Honours. On 12 March 1944 the Japanese launched an invasion of India aimed at Imphal, hundreds of miles to the north. General Renya Mutaguchi of the Burma Area Army announced the invasion of India was "The March into Delhi" as he expected the invasion to end with him marching in triumph into New Delhi. Slim knew from signals intelligence that the Japanese were going to invade in March 1944, but as Murray and Millet wrote  "...he had little choice, but to meet it with the forces on hand – the IV Corps of three Anglo-Indian divisions – or surrender his own plans to take the general offensive into Burma in 1944." Slim chose to fight a defensive campaign to break the Japanese before launching his offensive into Burma, believing that superior British tanks, logistics and air power would allow him to inflict a decisive defeat on Mutaguchi. However, the Japanese advanced more swiftly than Slim had expected up the mud roads of Burma into India, leading to a period of crisis as the fate of India hung in the balance. Slim airlifted two entire veteran divisions (5th and 7th Indian) from battle in the Arakan, straight into battle in the north. Desperate defensive actions were fought at places such as Imphal, Sangshak and Kohima, while the RAF and USAAF kept the forces supplied from the air. Slim ordered his men to hold their ground, forbade any retreat, and informed his men who were surrounded by the Japanese that supplies would be flown in from the air to allow them to hold out. Slim decided to send the IV Corps to relieve Imphal while gambling that the 5th Indian Division could hold out at Kohima, though Slim knew that if Kohima fell, then the Japanese could be able to sever the Assam railroad at Dimapur, which could cut off the British Fourteenth Army from its main supply line. The Battle of Kohima was a fiercely fought battle as Murray and Millet wrote: "Nowhere in World War II – even on the Eastern Front – did the combatants fight with more mindless savagery", but Kohima held. As late as 1 June 1944, Field Marshal Alan Brooke, the Chief of the Imperial General Staff, wrote in his diary that he saw "disaster staring us in the face" in Assam, but Slim was more confident, believing he could smash the Japanese attempt to take India. While the Japanese were able to advance and encircle the formations of British Fourteenth Army, they were unable to defeat those same forces or break out of the jungles along the Indian frontier. The Japanese advance stalled. The Japanese, who had a contempt for British and Indian troops based on their performance in 1941–42, refused to give up even after the monsoon started and large parts of their army were wrecked by conducting operations in impossible conditions. The initial Japanese plan was to capture Allied stocks of food, medicine and fuel to sustain their advance, but they failed to capture any stockpiles. As a result, their units took unsupportable casualties and were finally forced to retreat in total disorder in July 1944, leaving behind many dead from hunger and disease as well as their injured. Of the 150,000 Japanese soldiers who invaded India in March 1944, almost all of them were dead by July 1944 as Slim had inflicted the largest defeats that the Japanese had suffered up to this point in the war. The Indian Army remained loyal, and under its British officers fought much better than the Indian National Army. After Imphal and Kohima, the American historian Gerhard Weinberg noted for the Japanese their "... only hope in the area was propaganda by Bose, not much of a substitute for their lost Japanese 15th Army".

Unlike the Japanese, who killed their wounded, Slim went out of his way to ensure good medical care for his wounded and to evacuate his wounded by air to hospitals in India. Slim knew his men would fight better if they knew that they would receive the best possible medical care under the conditions if they were wounded. On 8 August 1944, Slim was promoted to lieutenant general, and, on 28 September 1944, he was appointed a Knight Commander of the Order of the Bath (KCB). In December 1944, during a ceremony at Imphal in front of the Scottish, Gurkha and Punjabi regiments, Slim and three of his corps commanders (Christison, Scoones and Stopford) were knighted by the viceroy Lord Wavell and invested with honours. Slim was presented with his insignia as KCB, and the others with their KBEs. Slim was also mentioned in despatches. By the end of 1944, the majority of the men serving in the British Fourteenth Army were in fact not British as of the 12 divisions that made up the British Fourteenth Army, 2 were British, 7 were Indian and 3 came from Britain's African colonies. In addition, there were six Chinese divisions, two regiments from the U.S. Army and various tribal militias made up of Shan, Chin, Naga, Kachin and Karen peoples raised by the OSS and the SOE fighting on the Allied side in Burma, requiring Slim to play the role of the diplomat as much as a general to hold these disparate forces made up of so many different peoples together.

In 1945, Slim launched an offensive into Burma, with supply lines stretching almost to the breaking point across hundreds of miles of trackless jungle. Slim employed Billy Williams and his corps of elephants, led by Bandoola, to build bridges and rescue refugees. He faced the same problems that the Japanese had faced in their failed 1944 offensive in the opposite direction. He made the supply of his armies the central issue in the plan of the campaign. The Chindwin River was spanned with the longest Bailey bridge in the world at the time. To distract the Japanese from his campaign in central and southern Burma, Slim ordered the Chinese in northern Burma to begin an offensive, which for a time led the Japanese to the erroneous conclusion that the main goal of the Allies was to open the Burma Road to China. Slim began his advance by sending two corps towards Mandalay and another corps along the coast towards Rangoon, but changed his plans when he learned from intelligence that the Japanese were planning on defending Mandalay from the eastern banks of the Irrawaddy River. Slim had one corps cross the Irrawaddy south of Mandalay at Meiktila while another corps staged a feint attack on Mandalay from the north to distract the Japanese from the main blow coming up from the south. The swift flowing Irrawaddy is a wider river than the Rhine, making it into a natural defensive barrier that the Japanese believed could halt the British advance. However, much of the countryside around the Irrawaddy consists of plains that favoured the offensive, and in his operations in the Irrawaddy river valley, Slim used combined arms offensives with artillery and tanks working closely with the infantry to bring down overwhelming firepower when the Japanese tried to block the British Fourteenth Army's advance. In March 1945, after crossing the Irrawaddy, the town of Meiktila was taken, followed by Burma's second city, Mandalay.

The Japanese garrison in Mandalay chose not to surrender, using the forts built by the British and the maze of pagodas in downtown Mandalay to fight to the death in an urban battle that destroyed much of the city, which finally fell to the British Fourteenth Army on 20 March 1945. Slim's plan was a masterpiece of operational art, and the capture of Meiktila left most of Japan's troops stranded in Burma without supplies. The Allies had reached the open plains of central Burma, sallying out and breaking Japanese attacking forces in isolation, maintaining the initiative at all times, backed up by air-land cooperation, including resupply by air and close air support, performed by both RAF and USAAF units. Slim followed up this victory by ordering his coastal corps to seize the mouth of the Irrawaddy where it flowed into the Bay of Bengal.

In combination with these attacks, Force 136 helped initiate a countrywide uprising of the Bamar people against the Japanese. In addition to fighting the allied advance south, the Japanese were faced with heavy attacks from behind their own lines. As he advanced into Burma, Slim discovered gruesome evidence of the nature of Japanese rule in Burma, finding in village after village, Burmese peasants who had been tied to trees and bayoneted to death as the Japanese preferred bayonet practice with people rather than sandbags as normally is the case. Toward the end of the campaign, the army raced south to capture Rangoon before the start of the monsoon. It was considered necessary to capture the port because of the length of the supply lines overland from India and the impossibility of supply by air or land during the monsoon. Rangoon was eventually taken by a combined attack from the land (Slim's army), the air (parachute operations south of the city) and a seaborne invasion. Also assisting in the capture of Rangoon was the Anti-Fascist People's Freedom League led by Thakin Soe, with Aung San (the future Prime Minister of Burma and father of Aung San Suu Kyi) as one of its military commanders.

As the Burma campaign came to an end, Slim was informed in May by Oliver Leese, the commander of Allied Land Forces South-East Asia (ALFSEA) that he would not be commanding Fourteenth Army in the forthcoming invasion planned for Malaya but would take command of the new Twelfth Army being formed to mop up in Burma. Slim refused the appointment, saying he would prefer to retire. As the news spread, Fourteenth Army fell into turmoil and Alan Brooke, the Chief of the Imperial General Staff, furious at not having been consulted by Leese, and Claude Auchinleck, the C-in-C India who was at the time in London, brought pressure to bear. The Supreme Allied Commander of the Southeast Asia Theatre, Louis Mountbatten was obliged to order Leese to undo the damage. On 1 July 1945, Slim was promoted to general and was informed that he was to succeed Leese as C-in-C ALFSEA. However, by the time he took up the post, having taken some leave, the war was at an end.

Relations with troops
Slim had an excellent relationship with his troops – the "Forgotten Army", as they called themselves. Slim had a close rapport with the officers and soldiers under his command, and always trusted his officers to make the correct decisions without referring to him. As Slim later wrote: "I was, like other generals before me, to be saved ... by the resourcefulness and the stubborn valour of my troops". Murray and Millet wrote that Slim's willingness to delegate authority down to his officers on the spot played a key role in sustaining his advance into Burma as officers did not have to wait for a decision from him.

In the aftermath of Kohima and Imphal, Slim inspected the battlefields, seeing the bodies of wounded Japanese soldiers who had been killed by their comrades as under the code of Bushido, it was considered shameful to surrender and it was the highest honour for a Japanese soldier to die for the Emperor, which graphically showed to Slim how far the Japanese were willing to take Bushido. Slim realized that Japanese logistics had broken down, but that Japanese soldiers, motivated by Bushido, were still prepared to fight to the death as the Japanese preferred to die for the Emperor rather than surrender, which led him to the conclusion that it was better to outflank and bypass the Japanese positions, leaving the main pockets of the Japanese to starve to death rather than engaging them in combat as much as possible. The Burma Area Army had about 100,000 men while the British Fourteenth Army had only about 21,000 men, but Slim decided with superior mobility backed by proper supply lines that he could defeat the Burma Area Army, whose logistics were poor. The superior Japanese numbers together with the fact that the Indian Army was an all-volunteer force, with only so many Indians willing to volunteer and the fact that sending fresh British troops to Burma was not a priority in London, made it imperative for Slim to save the lives of his men as much as possible. Slim was painfully aware that it would be difficult to replace whatever losses his men took, and had no intention of having his army being ground down by fighting the Japanese in every single place that they were. The differences between Slim, who was determined to save the lives of his men as much as possible versus the fanatical desire of Japanese officers motivated by Bushido to have every man under their command die for the Emperor, led Slim to estimate that for every man killed under his command, the Japanese lost a hundred men.

In Slim's book, Defeat into Victory, he tells of the malaria rates among his units being 70%, largely due to noncompliance by his soldiers with the foul-tasting mepacrine they refused to take. Slim did not blame his medics for this problem, but placed the responsibility on his officers. "Good doctors are no use without good discipline. More than half the battle against disease is fought not by the doctors, but by the regimental officers". After Slim dismissed a few officers for high unit malaria rates, the others realised he was serious and malaria treatment was enforced, dropping the rate to less than five per cent. The combat effectiveness of his army was thus greatly enhanced. This physical and mental turnaround in the army under him was a contributing factor to the eventual defeat of the Japanese in Burma.

George MacDonald Fraser, later author of The Flashman Papers series of novels, then a nineteen-year-old lance corporal, recalled:

and:

Post-war career

Initial retirement from the army
At the end of 1945 Slim returned to the UK. On 1 January 1946, he was made a Knight Grand Cross of the Order of the British Empire (GBE). and took the post of Commandant of the Imperial Defence College for its first course since 1939. On 7 February 1947 he was made an Aide-de-camp (ADC) to the King. At the end of his two-year appointment at the Imperial Defence College Slim retired as ADC and from the army on 11 May 1948. He had been approached by both India and Pakistan to become C-in-C of their respective armies post independence but refused and instead became Deputy Chairman of the Railway Executive.

Return to the army
However, in November 1948 the British Prime Minister Clement Attlee rejected the proposal by Viscount Montgomery that he should be succeeded as Chief of the Imperial General Staff (CIGS) by John Crocker and instead brought back Slim from retirement in the rank of field marshal in January 1949 with formal appointment to the Army Council from 1 January 1949. Slim thus became the first Indian Army officer to become CIGS.

On 2 January 1950, he was promoted to Knight Grand Cross of the Order of the Bath (GCB) and later that year was made a Chief Commander of the Legion of Merit by the United States having previously, in 1948, been awarded the lower ranking Commander of the Legion of Merit. On 1 November 1952, he relinquished the position of Chief of the Imperial General Staff.

Governor-General of Australia
On 10 December 1952 Slim was made a Knight Grand Cross of the Order of St Michael and St George (GCMG) on his appointment as Governor-General of Australia which post he took up on 8 May 1953. On 2 January 1953, he was appointed a Knight of the Order of St. John (KStJ). Slim was a popular choice for Governor-General since he was an authentic war hero who had fought alongside Australians at Gallipoli and in the Middle East. In 1954 he was able to welcome Queen Elizabeth II on the first visit by a reigning monarch to Australia. For his services to the Queen during the tour, he was appointed a Knight Grand Cross of the Royal Victorian Order (GCVO) on 27 April 1954. Liberal leader Robert Menzies held office throughout Slim's time in Australia. His Official Secretary throughout his term was Murray Tyrrell.

Retirement
In 1959, Slim retired and returned to Britain, where he published his memoirs, Unofficial History. He had already published his personal narrative of the Burma Campaign, Defeat into Victory, in 1956, which has never been out of print, and in which he candidly talked about his mistakes and the lessons he learned. On 24 April 1959, he was appointed a Knight Companion of the Order of the Garter (KG). On 15 July 1960, he was created "Viscount Slim of Yarralumla in the Capital Territory of Australia and of Bishopston in the City and County of Bristol".

After a successful further career on the boards of major UK companies, he was appointed Constable and Governor of Windsor Castle on 18 June 1964. He died in London on 14 December 1970, aged 79 and was given a full military funeral at St. George's Chapel, Windsor and was afterwards cremated.

Allegations
During his tenure as Governor-General of Australia, Slim was patron of the Fairbridge Farm school child migration to Australia. Forty-seven years later, in 2007, and thirty-seven years after Slim's death, three former child migrants alleged Slim sexually assaulted them during visits to Fairbridge Farm. The allegations were dismissed at the time both by those who had served under Slim in the army and by his son. The allegations were aired again on ABC television in the programme The Long Journey Home, broadcast on 17 November 2009. On 27 February 2017, the ABC again aired allegations against Slim and Fairbridge Farm.

Eponyms
 William Slim Drive, in the Canberra district of Belconnen, was named after him. Despite an independent review commissioned by the Australian Capital Territory Government, which returned no recommendation to change the road name, in 2019, the then ACT Planning Minister Mick Gentleman, announced that, after considering allegations made, submissions to the Royal Commission into Institutional Responses to Child Sexual Abuse and submissions by the Slim family, he would change the road's name and in 2021 the name was changed.
 The Slim Officers' Mess at the Royal Military Academy Sandhurst is named after him and was opened in August 2004 by his son.
 On 7 September 2008, a plaque in Slim's memory, and those who served with him, was unveiled at the Cenotaph in his native Bristol. Fr Robert King of the Diocese of Clifton was joined by faith leaders from the Hindu, Muslim and Sikh communities at the ceremony, which was led by the Lord Mayor of Bristol's chaplain, the Rev. Prebendary Harold Clarke.
 The road Viscount Slim Avenue, in Whyalla, South Australia is named after him.
 The Slim Building at the Cranfield University Shrivenham Campus is named after him.
 The Slim School was a British Families Education Service co-educational secondary school located in the Cameron Highlands, Malaya, and was named after him.

Historical assessment

Lieutenant General Sir John Kiszely has recommended Slim's memoirs (Defeat into Victory) (1956) in which Slim candidly talked about his own mistakes in the Burma campaign. The book has never been out of print.  Kiszely has described Slim as "perhaps the Greatest Commander of the 20th Century" and has commenting on Slim's "self-deprecating style" Military historian Max Hastings stated:

In contrast to almost every other outstanding commander of the war, Slim was a disarmingly normal human being, possessed of notable self-knowledge. He was without pretension, devoted to his wife, Aileen, their family and the Indian Army. His calm, robust style of leadership and concern for the interests of his men won the admiration of all who served under him ... His blunt honesty, lack of bombast and unwillingness to play courtier did him few favours in the corridors of power. Only his soldiers never wavered in their devotion.

The spirit of comradeship Slim created within Fourteenth Army lived on after the war in the Burma Star Association, of which Slim was a co-founder and first President.

A statue of Slim on Whitehall, outside the Ministry of Defence, was unveiled by Queen Elizabeth II in 1990. Designed by Ivor Roberts-Jones, the statue is one of three British Second World War military leaders (the others being Alan Brooke and Bernard Montgomery).

Slim's papers were collected by his biographer, Ronald Lewin, and given to the Churchill Archives Centre by Slim's wife, Aileen, Viscountess Slim, and son, John Slim, 2nd Viscount Slim, and other donors, 1977–2001. Lewin's biography, Slim: The Standardbearer, was awarded the 1977 WH Smith Literary Award.

Arms

List of honours

Bibliography
 Defeat into Victory by Field Marshal Sir William Slim; Originally published 1956. More recent editions are NY: Buccaneer Books , Cooper Square Press ; London: Cassell , Pan .
 Other publications include Courage and Other Broadcasts (1957); and Unofficial History (1959).

Footnotes

References

  Also 1999 Wordsworth Military Library softcover reprint:  .

  (Stone & Stone, 1 August 1997. Note: The original edition contains more detailed description of operations than does the 1961 edition.)

Further reading

External links

 : "The joint winners of Britain's Greatest General, as decided by the audience at [the National Army Museum] celebrity speaker event on Saturday 9 April 2011, were William Slim and The Duke of Wellington".
 Generals of World War II
 Indian Army Officers 1939–1945

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1891 births
1970 deaths
Military personnel from Bristol
Governors-General of Australia
Royal Warwickshire Fusiliers officers
West India Regiment officers
British Army personnel of World War I
Indian Army generals of World War II
British field marshals
English Roman Catholics
Chief Commanders of the Legion of Merit
People educated at King Edward's School, Birmingham
People educated at St Philip's School
Chiefs of the Imperial General Staff
People associated with the University of Birmingham
Burma in World War II
Graduates of the Royal College of Defence Studies
Graduates of the Staff College, Quetta
Viscounts created by Elizabeth II
Indian Army personnel of World War I
Commandants of the Senior Officers' School, Belgaum
Academics of the Staff College, Camberley
Pakistan Command and Staff College alumni

William Slim, 1st Viscount Slim


Rauno "Rane" Korpi (born June 25, 1951 in Tampere, Finland) is a Finnish ice hockey coach. He coached Tappara to three consecutive Finnish Championships in SM-liiga during the years 1986-1988. He has also won one additional Finnish Championship Gold, one Silver medal and one Bronze medal. Korpi has been chosen three times as The Coach of the Year in Finland. Other teams coached by Korpi include TuTo, EC KAC, and EV Zug.

Korpi has also coached Finland's national ice hockey team during the years 1986-1987. He has coached Women's national teams and Junior national teams as well.

Korpi is the father of Kiira Korpi, Olympic-level figure skater and winner of European Championships Bronze medal.

References

1951 births
Living people
Finnish ice hockey players
Finland men's national ice hockey team coaches
Ice hockey people from Tampere

Rauno Korpi


The Philippines' Department of Social Welfare and Development (, abbreviated as DSWD) is the executive department of the Philippine Government responsible for the protection of the social welfare of rights of Filipinos and to promote the social development.

History

In 1915, the Public Welfare Board (PWB) was created and was tasked to study, coordinate and regulate all government and private entities engaged in social services. In 1921, the PWB was abolished and replaced by the Bureau of Public Welfare under the Department of Public Instruction.

On November 1, 1939, Commonwealth Act No. 439 created the Department of Health and Public Welfare and in 1941, the Bureau of Public Welfare officially became a part of the Department of Health and Public Welfare. In addition to coordinating services of all public and private social welfare institutions, the Bureau also managed all public child-caring institutions and the provision of child welfare services.

In 1947, President Manuel Roxas abolished the Bureau of Public Welfare and created the Social Welfare Commission, under the Office of the President, in its place.

In 1968, Republic Act 5416, known as the Social Welfare Act of 1968, created the Department of Social Welfare, placing it under the executive branch of government. In 1976, the Department of Social Welfare was renamed Department of Social Services and Development (DSSD) through Presidential Decree No. 994. This was signed into law by President Ferdinand E. Marcos and gave the department an accurate institutional identity. On June 2, 1978, the DSSD was renamed Ministry of Social Services and Development (MSSD) in line with the change in the form of government.

In 1987, the MSSD was reorganized and renamed the Department of Social Welfare and Development (DSWD) through Executive Order 123, which was signed by President Corazon C. Aquino. Executive Order No. 292, also known as the Revised Administration Code of 1987, established the name, organizational structure, and functional areas of responsibility of DSWD and further defined its statutory authority.

In 1991, the passage of Republic Act No. 7160 otherwise known as the Local Government Code of 1991 affected the devolution of DSWD basic services to local government units.

Organizational structure
The department is headed by the Secretary of Social Welfare and Development (Philippines), with the following eleven undersecretaries and nineteen assistant secretaries:
Undersecretary for Social Welfare and Development
Undersecretary for General Administration and Support Services Group
Undersecretary for Operations Group
Undersecretary for Disaster Response Management Group
Undersecretary for Policy and Plans Group
Undersecretary for Standards and Capacity Building
Undersecretary for National Household Targeting System and Pantawid Pamilyang Pilipino Program
Undersecretary for Special Concerns
Undersecretary for Innovations
Undersecretary for Inclusive and Sustainable Peace
Undersecretary for Legislative Coordination and External Affairs
Assistant Secretary for Strategic Communications
Assistant Secretary for Policy and Plans
Assistant Secretary for Legislative Affairs under PPG
Assistant Secretary for External Assistance and Development under PPG
Assistant Secretary for Administration under GASSG
Assistant Secretary for Finance under GASSG
Assistant Secretary for SCBG
Assistant Secretary for DRMG
Assistant Secretary for Specialized Programs under Operations Group
Assistant Secretary for Statutory Programs under Operations Group
Assistant Secretary for Regional Operations under Operations Group
Assistant Secretary for the National Household Targeting System for Poverty Reduction
Assistant Secretary for the Pantawid Pamilyang Pilipino Program
Assistant Secretary for Inclusive-Sustainable Peace and Special Concerns
Assistant Secretary for Community Engagement
Assistant Secretary for Innovations
Assistant Secretary for Partnerships Building and Resource Mobilization under the Office of the Secretary
Assistant Secretary for International Affairs, and Attached and Supervised Agencies under the Office of the Secretary
Assistant Secretary/Chief Information Officer under the Office of the Secretary
Assistant Secretary/Special Assistant to the Secretary for Centers and Residential Care Facilities

Attached Agencies
 Council for the Welfare of Children
 National Authority for Child Care (NACC)
 Juvenile Justice and Welfare Council
 National Council on Disability Affairs

Supervised Agencies
 National Anti-Poverty Commission
 National Commission on Indigenous Peoples
 Presidential Commission for the Urban Poor

Programs and Services

The Pantawid Pamilyang Pilipino Program
The Pantawid Pamilyang Pilipino Program or "4Ps" (conditional cash transfer) is a human development program that invests in the health and education of poor families, primarily those with children aged 0–18.

Kalahi CIDSS – NCDDP
The Kapit-Bisig Laban sa Kahirapan – Comprehensive and Integrated Delivery of Social Services – National Community-Driven Development Program (Kalahi CIDSS–NCDDP) is the community-driven development program of the Philippine Government implemented through the Department of Social Welfare and Development. Supplemented by the government of the Philippines.

Sustainable Livelihood Program
The Sustainable Livelihood Program (SLP) is a community-based capacity building effort that seeks to improve the program participants' socio-economic status through two tracks: Micro-enterprise Development and Employment Facilitation.

Listahanan
An information management system that identifies who and where the poor are in the country. It is being operated by the National Household Targeting System for Poverty Reduction (NHTS-PR).

Supplemental Feeding Program
Provision of food in addition to the regular meals, to target children as part of the DSWD's ECCD program of the government.

Disaster Response Operations
Life-saving emergency relief and long-term response.

RRPTP
Recovery and Reintegration Program for Trafficked Persons (RRTP) is a comprehensive package of programs and services, enhancing the psychosocial and economic needs of the beneficiaries.

PAMANA
Payapa at Masaganang Pamayanan (PAMANA) aims to improve access of poor communities to basic social services and promote responsive governance.

Protective Services Program
Assistance to Individuals In Crisis Situations (AICS) and Assistance to Communities in Needs (ACN)
Provides a range of interventions to individuals, families, and communities in crisis or difficult situations and vulnerable or disaster-affected communities.

ISWSFN
International Social Welfare Services for Filipino Nationals is a program for migrant Filipinos and other overseas Filipino nationals who are in crisis situation and in need of special protection are encouraged to seek assistance in the Philippine Embassies in their countries of destination.

Center & Residential Care Facilities
Services rendered in facilities 24-hour that provide alternative family care arrangement to poor, vulnerable and disadvantaged individuals or families in crisis.

Adoption and Foster Care
The act of adoption, of permanently placing a minor with a parent or parents other than the birth parents in the Philippines.

Gender and Development
Gender is about relations—between men and women, women and women, also between men and men and boys and girls. The GAD as perspective recognizes that gender concerns cut across all areas of development and therefore gender must influence government when it plans, budget for, implements, monitors and evaluates policies, programs and projects for development.

BUB
Pilot tested in 2013 and now on its 3rd cycle, the Bottom-Up Budgeting (BUB) Process is proposed to ensure implementation of priority poverty reduction projects.

Secretaries

References

3. DSWD

External links

Department of Social Welfare and Development
Social Welfare and Development
Philippines
Welfare in the Philippines
Philippines, Social Welfare and Development
1939 establishments in the Philippines


Evolutionary developmental biology (informally, evo-devo) is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.

The field grew from 19th-century beginnings, where embryology faced a mystery: zoologists did not know how embryonic development was controlled at the molecular level. Charles Darwin noted that having similar embryos implied common ancestry, but little progress was made until the 1970s. Then, recombinant DNA technology at last brought embryology together with molecular genetics. A key early discovery was of homeotic genes that regulate development in a wide range of eukaryotes.

The field is composed of multiple core evolutionary concepts. One is deep homology, the finding that dissimilar organs such as the eyes of insects, vertebrates and cephalopod molluscs, long thought to have evolved separately, are controlled by similar genes such as pax-6, from the evo-devo gene toolkit. These genes are ancient, being highly conserved among phyla; they generate the patterns in time and space which shape the embryo, and ultimately form the body plan of the organism. Another is that species do not differ much in their structural genes, such as those coding for enzymes; what does differ is the way that gene expression is regulated by the toolkit genes. These genes are reused, unchanged, many times in different parts of the embryo and at different stages of development, forming a complex cascade of control, switching other regulatory genes as well as structural genes on and off in a precise pattern. This multiple pleiotropic reuse explains why these genes are highly conserved, as any change would have many adverse consequences which natural selection would oppose.

New morphological features and ultimately new species are produced by variations in the toolkit, either when genes are expressed in a new pattern, or when toolkit genes acquire additional functions. Another possibility is the neo-Lamarckian theory that epigenetic changes are later consolidated at gene level, something that may have been important early in the history of multicellular life.

History

Early theories

Philosophers began to think about how animals acquired form in the womb in classical antiquity. Aristotle asserts in his Physics treatise that according to Empedocles, order "spontaneously" appears in the developing embryo. In his The Parts of Animals treatise, he argues that Empedocles' theory was wrong. In Aristotle's account, Empedocles stated that the vertebral column is divided into vertebrae because, as it happens, the embryo twists about and snaps the column into pieces. Aristotle argues instead that the process has a predefined goal: that the "seed" that develops into the embryo began with an inbuilt "potential" to become specific body parts, such as vertebrae. Further, each sort of animal gives rise to animals of its own kind: humans only have human babies.

Recapitulation

A recapitulation theory of evolutionary development was proposed by Étienne Serres in 1824–26, echoing the 1808 ideas of Johann Friedrich Meckel. They argued that the embryos of 'higher' animals went through or recapitulated a series of stages, each of which resembled an animal lower down the great chain of being. For example, the brain of a human embryo looked first like that of a fish, then in turn like that of a reptile, bird, and mammal before becoming clearly human. The embryologist Karl Ernst von Baer opposed this, arguing in 1828 that there was no linear sequence as in the great chain of being, based on a single body plan, but a process of epigenesis in which structures differentiate. Von Baer instead recognized four distinct animal body plans: radiate, like starfish; molluscan, like clams; articulate, like lobsters; and vertebrate, like fish. Zoologists then largely abandoned recapitulation, though Ernst Haeckel revived it in 1866.

Evolutionary morphology

From the early 19th century through most of the 20th century, embryology faced a mystery. Animals were seen to develop into adults of widely differing body plan, often through similar stages, from the egg, but zoologists knew almost nothing about how embryonic development was controlled at the molecular level, and therefore equally little about how developmental processes had evolved. Charles Darwin argued that a shared embryonic structure implied a common ancestor. For example, Darwin cited in his 1859 book On the Origin of Species the shrimp-like larva of the barnacle, whose sessile adults looked nothing like other arthropods; Linnaeus and Cuvier had classified them as molluscs. Darwin also noted Alexander Kowalevsky's finding that the tunicate, too, was not a mollusc, but in its larval stage had a notochord and pharyngeal slits which developed from the same germ layers as the equivalent structures in vertebrates, and should therefore be grouped with them as chordates.

19th century zoology thus converted embryology into an evolutionary science, connecting phylogeny with homologies between the germ layers of embryos. Zoologists including Fritz Müller proposed the use of embryology to discover phylogenetic relationships between taxa. Müller demonstrated that crustaceans shared the Nauplius larva, identifying several parasitic species that had not been recognized as crustaceans. Müller also recognized that natural selection must act on larvae, just as it does on adults, giving the lie to recapitulation, which would require larval forms to be shielded from natural selection. Two of Haeckel's other ideas about the evolution of development have fared better than recapitulation: he argued in the 1870s that changes in the timing (heterochrony) and changes in the positioning within the body (heterotopy) of aspects of embryonic development would drive evolution by changing the shape of a descendant's body compared to an ancestor's. It took a century before these ideas were shown to be correct.

In 1917, D'Arcy Thompson wrote a book on the shapes of animals, showing with simple mathematics how small changes to parameters, such as the angles of a gastropod's spiral shell, can radically alter an animal's form, though he preferred a mechanical to evolutionary explanation. But without molecular evidence, progress stalled.

In 1952, Alan Turing published his paper "The Chemical Basis of Morphogenesis", on the development of patterns in animals' bodies. He suggested that morphogenesis could be explained by a reaction–diffusion system, a system of reacting chemicals able to diffuse through the body. He modelled catalysed chemical reactions using partial differential equations, showing that patterns emerged when the chemical reaction produced both a catalyst (A) and an inhibitor (B) that slowed down production of A. If A and B then diffused at different rates, A dominated in some places, and B in others. The Russian biochemist Boris Belousov had run experiments with similar results, but was unable to publish them because scientists thought at that time that creating visible order violated the second law of thermodynamics.

The modern synthesis of the early 20th century

In the so-called modern synthesis of the early 20th century, between 1918 and 1930 Ronald Fisher brought together Darwin's theory of evolution, with its insistence on natural selection, heredity, and variation, and Gregor Mendel's laws of genetics into a coherent structure for evolutionary biology. Biologists assumed that an organism was a straightforward reflection of its component genes: the genes coded for proteins, which built the organism's body. Biochemical pathways (and, they supposed, new species) evolved through mutations in these genes. It was a simple, clear and nearly comprehensive picture: but it did not explain embryology. Sean B. Carroll has commented that had evo-devo's insights been available, embryology would certainly have played a central role in the synthesis.

The evolutionary embryologist Gavin de Beer anticipated evolutionary developmental biology in his 1930 book Embryos and Ancestors, by showing that evolution could occur by heterochrony, such as in the retention of juvenile features in the adult. This, de Beer argued, could cause apparently sudden changes in the fossil record, since embryos fossilise poorly. As the gaps in the fossil record had been used as an argument against Darwin's gradualist evolution, de Beer's explanation supported the Darwinian position. However, despite de Beer, the modern synthesis largely ignored embryonic development to explain the form of organisms, since population genetics appeared to be an adequate explanation of how forms evolved.

The lac operon

In 1961, Jacques Monod, Jean-Pierre Changeux and François Jacob discovered the lac operon in the bacterium Escherichia coli. It was a cluster of genes, arranged in a feedback control loop so that its products would only be made when "switched on" by an environmental stimulus. One of these products was an enzyme that splits a sugar, lactose; and lactose itself was the stimulus that switched the genes on. This was a revelation, as it showed for the first time that genes, even in organisms as small as a bacterium, are subject to precise control. The implication was that many other genes were also elaborately regulated.

The birth of evo-devo and a second synthesis
In 1977, a revolution in thinking about evolution and developmental biology began, with the arrival of recombinant DNA technology in genetics, the book Ontogeny and Phylogeny by Stephen J. Gould and the paper "Evolution and Tinkering" by François Jacob. Gould laid to rest Haeckel's interpretation of evolutionary embryology, while Jacob set out an alternative theory.
This led to a second synthesis, at last including embryology as well as molecular genetics, phylogeny, and evolutionary biology to form evo-devo. In 1978, Edward B. Lewis discovered homeotic genes that regulate embryonic development in Drosophila fruit flies, which like all insects are arthropods, one of the major phyla of invertebrate animals.
Bill McGinnis quickly discovered homeotic gene sequences, homeoboxes, in animals in other phyla, in vertebrates such as frogs, birds, and mammals; they were later also found in fungi such as yeasts, and in plants. There were evidently strong similarities in the genes that controlled development across all the eukaryotes.
In 1980, Christiane Nüsslein-Volhard and Eric Wieschaus described gap genes which help to create the segmentation pattern in fruit fly embryos; they and Lewis won a Nobel Prize for their work in 1995.

Later, more specific similarities were discovered: for example, the Distal-less gene was found in 1989 to be involved in the development of appendages or limbs in fruit flies, the fins of fish, the wings of chickens, the parapodia of marine annelid worms, the ampullae and siphons of tunicates, and the tube feet of sea urchins. It was evident that the gene must be ancient, dating back to the last common ancestor of bilateral animals (before the Ediacaran Period, which began some 635 million years ago). Evo-devo had started to uncover the ways that all animal bodies were built during development.

The control of body structure

Deep homology

Roughly spherical eggs of different animals give rise to unique morphologies, from jellyfish to lobsters, butterflies to elephants. Many of these organisms share the same structural genes for bodybuilding proteins like collagen and enzymes, but biologists had expected that each group of animals would have its own rules of development. The surprise of evo-devo is that the shaping of bodies is controlled by a rather small percentage of genes, and that these regulatory genes are ancient, shared by all animals. The giraffe does not have a gene for a long neck, any more than the elephant has a gene for a big body. Their bodies are patterned by a system of switching which causes development of different features to begin earlier or later, to occur in this or that part of the embryo, and to continue for more or less time.

The puzzle of how embryonic development was controlled began to be solved using the fruit fly Drosophila melanogaster as a model organism. The step-by-step control of its embryogenesis was visualized by attaching fluorescent dyes of different colours to specific types of protein made by genes expressed in the embryo. A dye such as green fluorescent protein, originally from a jellyfish, was typically attached to an antibody specific to a fruit fly protein, forming a precise indicator of where and when that protein appeared in the living embryo.

Using such a technique, in 1994 Walter Gehring found that the pax-6 gene, vital for forming the eyes of fruit flies, exactly matches an eye-forming gene in mice and humans. The same gene was quickly found in many other groups of animals, such as squid, a cephalopod mollusc. Biologists including Ernst Mayr had believed that eyes had arisen in the animal kingdom at least 40 times, as the anatomy of different types of eye varies widely. For example, the fruit fly's compound eye is made of hundreds of small lensed structures (ommatidia); the human eye has a blind spot where the optic nerve enters the eye, and the nerve fibres run over the surface of the retina, so light has to pass through a layer of nerve fibres before reaching the detector cells in the retina, so the structure is effectively "upside-down"; in contrast, the cephalopod eye has the retina, then a layer of nerve fibres, then the wall of the eye "the right way around". The evidence of pax-6, however, was that the same genes controlled the development of the eyes of all these animals, suggesting that they all evolved from a common ancestor. Ancient genes had been conserved through millions of years of evolution to create dissimilar structures for similar functions, demonstrating deep homology between structures once thought to be purely analogous. This notion was later extended to the evolution of embryogenesis and has caused a radical revision of the meaning of homology in evolutionary biology.

Gene toolkit

A small fraction of the genes in an organism's genome control the organism's development. These genes are called the developmental-genetic toolkit. They are highly conserved among phyla, meaning that they are ancient and very similar in widely separated groups of animals. Differences in deployment of toolkit genes affect the body plan and the number, identity, and pattern of body parts. Most toolkit genes are parts of signalling pathways: they encode transcription factors, cell adhesion proteins, cell surface receptor proteins and signalling ligands that bind to them, and secreted morphogens that diffuse through the embryo. All of these help to define the fate of undifferentiated cells in the embryo. Together, they generate the patterns in time and space which shape the embryo, and ultimately form the body plan of the organism. Among the most important toolkit genes are the Hox genes. These transcription factors contain the homeobox protein-binding DNA motif, also found in other toolkit genes, and create the basic pattern of the body along its front-to-back axis.
Hox genes determine where repeating parts, such as the many vertebrae of snakes, will grow in a developing embryo or larva. Pax-6, already mentioned, is a classic toolkit gene. Although other toolkit genes are involved in establishing the plant bodyplan, homeobox genes are also found in plants, implying they are common to all eukaryotes.

The embryo's regulatory networks

The protein products of the regulatory toolkit are reused not by duplication and modification, but by a complex mosaic of pleiotropy, being applied unchanged in many independent developmental processes, giving pattern to many dissimilar body structures. The loci of these pleiotropic toolkit genes have large, complicated and modular cis-regulatory elements. For example, while a non-pleiotropic rhodopsin gene in the fruit fly has a cis-regulatory element just a few hundred base pairs long, the pleiotropic eyeless cis-regulatory region contains 6 cis-regulatory elements in over 7000 base pairs. The regulatory networks involved are often very large. Each regulatory protein controls "scores to hundreds" of cis-regulatory elements. For instance, 67 fruit fly transcription factors controlled on average 124 target genes each. All this complexity enables genes involved in the development of the embryo to be switched on and off at exactly the right times and in exactly the right places. Some of these genes are structural, directly forming enzymes, tissues and organs of the embryo. But many others are themselves regulatory genes, so what is switched on is often a precisely-timed cascade of switching, involving turning on one developmental process after another in the developing embryo.

Such a cascading regulatory network has been studied in detail in the development of the fruit fly embryo. The young embryo is oval in shape, like a rugby ball. A small number of genes produce messenger RNAs that set up concentration gradients along the long axis of the embryo. In the early embryo, the bicoid and hunchback genes are at high concentration near the anterior end, and give pattern to the future head and thorax; the caudal and nanos genes are at high concentration near the posterior end, and give pattern to the hindmost abdominal segments. The effects of these genes interact; for instance, the Bicoid protein blocks the translation of caudal messenger RNA, so the Caudal protein concentration becomes low at the anterior end. Caudal later switches on genes which create the fly's hindmost segments, but only at the posterior end where it is most concentrated.

The Bicoid, Hunchback and Caudal proteins in turn regulate the transcription of gap genes such as giant, knirps, Krüppel, and tailless in a striped pattern, creating the first level of structures that will become segments. The proteins from these in turn control the pair-rule genes, which in the next stage set up 7 bands across the embryo's long axis. Finally, the segment polarity genes such as engrailed split each of the 7 bands into two, creating 14 future segments.

This process explains the accurate conservation of toolkit gene sequences, which has resulted in deep homology and functional equivalence of toolkit proteins in dissimilar animals (seen, for example, when a mouse protein controls fruit fly development). The interactions of transcription factors and cis-regulatory elements, or of signalling proteins and receptors, become locked in through multiple usages, making almost any mutation deleterious and hence eliminated by natural selection.

The mechanism that sets up every animal's front-back axis is the same, implying a common ancestor. There is a similar mechanism for the back-belly axis for bilaterian animals, but it is reversed between arthropods and vertebrates. Another process, gastrulation of the embryo, is driven by Myosin II molecular motors, which are not conserved across species. The process may have been started by movements of sea water in the environment, later replaced by the evolution of tissue movements in the embryo.

The origins of novelty

Among the more surprising and, perhaps, counterintuitive (from a neo-Darwinian viewpoint) results of recent research in evolutionary developmental biology is that the diversity of body plans and morphology in organisms across many phyla are not necessarily reflected in diversity at the level of the sequences of genes, including those of the developmental genetic toolkit and other genes involved in development. Indeed, as John Gerhart and Marc Kirschner have noted, there is an apparent paradox: "where we most expect to find variation, we find conservation, a lack of change". So, if the observed morphological novelty between different clades does not come from changes in gene sequences (such as by mutation), where does it come from? Novelty may arise by mutation-driven changes in gene regulation.

Variations in the toolkit

Variations in the toolkit may have produced a large part of the morphological evolution of animals. The toolkit can drive evolution in two ways. A toolkit gene can be expressed in a different pattern, as when the beak of Darwin's large ground-finch was enlarged by the BMP gene, or when snakes lost their legs as distal-less became under-expressed or not expressed at all in the places where other reptiles continued to form their limbs. Or, a toolkit gene can acquire a new function, as seen in the many functions of that same gene, distal-less, which controls such diverse structures as the mandible in vertebrates, legs and antennae in the fruit fly, and eyespot pattern in butterfly wings. Given that small changes in toolbox genes can cause significant changes in body structures, they have often enabled the same function convergently or in parallel. distal-less generates wing patterns in the butterflies Heliconius erato and Heliconius melpomene, which are Müllerian mimics. In so-called facilitated variation, their wing patterns arose in different evolutionary events, but are controlled by the same genes. Developmental changes can contribute directly to speciation.

Consolidation of epigenetic changes

Evolutionary innovation may sometimes begin in Lamarckian style with epigenetic alterations of gene regulation or phenotype generation, subsequently consolidated by changes at the gene level. Epigenetic changes include modification of DNA by reversible methylation, as well as nonprogrammed remoulding of the organism by physical and other environmental effects due to the inherent plasticity of developmental mechanisms. The biologists Stuart A. Newman and Gerd B. Müller have suggested that organisms early in the history of multicellular life were more susceptible to this second category of epigenetic determination than are modern organisms, providing a basis for early macroevolutionary changes.

Developmental bias

Development in specific lineages can be biased either positively, towards a given trajectory or phenotype, or negatively, away from producing certain types of change; either may be absolute (the change is always or never produced) or relative. Evidence for any such direction in evolution is however hard to acquire and can also result from developmental constraints that limit diversification. For example, in the gastropods, the snail-type shell is always built as a tube that grows both in length and in diameter; selection has created a wide variety of shell shapes such as flat spirals, cowries and tall turret spirals within these constraints. Among the centipedes, the Lithobiomorpha always have 15 trunk segments as adults, probably the result of a developmental bias towards an odd number of trunk segments. Another centipede order, the Geophilomorpha, the number of segments varies in different species between 27 and 191, but the number is always odd, making this an absolute constraint; almost all the odd numbers in that range are occupied by one or another species.

Ecological evolutionary developmental biology

Ecological evolutionary developmental biology integrates research from developmental biology and ecology to examine their relationship with evolutionary theory. Researchers study concepts and mechanisms such as developmental plasticity, epigenetic inheritance, genetic assimilation, niche construction and symbiosis.

See also

 Arthropod head problem
 Cell signaling
 Evolution & Development (journal)
 Human evolutionary developmental biology
 Just So Stories (as seen by evolutionary developmental biologists)
 Plant evolutionary developmental biology
 Recapitulation theory

Notes

References

Sources

External links

Subfields of evolutionary biology
Developmental biology
Extended evolutionary synthesis

Evolutionary developmental biology


Thutmose I (also known as Thothmes, Thutmosis or Tuthmosis I, meaning Thoth-Born) was the third Pharaoh of the 18th dynasty of Ancient Egypt. He became pharaoh after the death of Amenhotep I. During his reign, he sent his armies into the Levant and Nubia, pushing the borders of Egypt further than ever before. He built many temples in Egypt and a tomb for himself in the Valley of the Kings. He is the first king known to have done this, though Amenhotep I may have done it first. He was succeeded by his son Thutmose II, and then his daughter, Hatshepsut. He ruled from 1506 to 1493 BC. Some scholars give his dates from 1526 BC to 1513 BC. This depends on where the astronomical observations used to calculate ancient Egyptian dates were made. If the observations were from the city of Memphis it would give the dates 20 years earlier than if it was Thebes.

Family
Thutmose's father is unknown, but it may have been Amenhotep I. His mother, Senseneb, was not from a royal family and may have been a lesser wife or concubine of Amenhotep. Thutmoses's wife, Queen Ahmose, the Great Royal Wife, was probably the daughter of Ahmose I and the sister of Amenhotep I. There is some doubt, as she was never called a "king's daughter." Some historians believe that she was Thutmose's own sister. Thutmose's first son with Ahmose was Amenmose. He was born long before Thutmose's coronation. He can be seen on a stela from about 1522 BC hunting near Memphis. He became the commander of the army. Amenmose died about the same time as Thutmose. Thutmose had another son, Wadjmose, and two daughters, Hatshepsut and Nefrubity, by Ahmose. Wadjmose died before his father, and Nefrubity died as an infant. Thutmose had a son by another wife, Mutnofret. This son, married to Thutmose's daughter Hatshepsut, succeeded him as Thutmose II. Hatshepsut said that Thutmose gave the kingship to them both. However, this is considered to be propaganda by Hatshepsut's supporters to support her claim to the throne when she later took over power.

Dates and length of reign
During the reign of Amenhotep I, the Egyptians recorded an astronomical event. This was the star Sothis appearing above the horizon at dawn. This is called a heliacal rising. Modern scientists believe that Sothis is the star that we call Sirius. If this observation was made at Thebes, it took place in 1517 BC. From this date, the date of Amenhotep's death, and Thutmose's coronation can be calculated to 1506 BC. But a few historians believe that the observations were made at either Heliopolis or Memphis. This would mean that Thutmose would have been crowned in 1526 BC.

An Egyptian priest, Manetho, writing in the third century BC, said that Thutmose I's reign lasted 12 Years and 9 Months (or 13 Years). This data is supported by two dated inscriptions from Years 8 and 9 of his reign with Thutmoses's cartouche found on a stone block in Karnak. Accordingly, Thutmose is usually said to have ruled from 1506 BC to 1493 BC, but some scholars date him from 1526 BC to 1513 BC

Military achievements

At the time Thutmose became king, Nubia rebelled against Egyptian rule. Thutmose travelled up the Nile with his army and fought in the battle, personally killing the Nubian king. He returned to Thebes with the Nubian king's body hung from the front of his ship. In the third year of his reign he led another expedition against Nubia. He ordered the canal at the first cataract to be made deeper to make it easier to get his ships upstream. The canal had been first built by Sesostris III of the 12th Dynasty. This helped make Nubia part of the Egyptian empire. This expedition is mentioned in two separate inscriptions by the king's son Thure:

In the second year of Thutmose's reign, a stele at Tombos records that he built a fortress there. This is near the third cataract. Before this the Egyptian army had stopped at Buhen, at the second cataract. This stele says that he already fought in Syria. His Syrian campaign may be placed at the beginning of the second year of his rule. The Syrian campaign was the farthest north any Egyptian ruler had ever campaigned. Although it has not been found in modern times, he set up a stele when he crossed the Euphrates River. During this campaign, the Syrian princes declared loyalty to Thutmose. However, after he went back to Egypt, they stopped tribute and began building defences against future attacks. Thutmose celebrated his victories with an elephant hunt in the area of Niy, near Apamea in Syria. He returned to Egypt with strange tales of the Euphrates, "that inverted water which flows upstream when it ought to be flowing downstream." The Euphrates was the first major river which the Egyptians had seen which flowed from the north. This would be downstream on the Nile, instead of the south which was upstream on the Nile. The river became known in Egypt as the "inverted water."

Nubia rebelled against Egyptian rule again in his fourth year. He sent his army even farther south to stop it. An inscription from his reign has been found as far south as Kurgus, which was south of the fourth cataract. During his reign, he began a number of projects which effectively ended Nubian independence for the next 500 years. He enlarged a temple to Sesostris III and Khnum, opposite the Nile from Semna. There are also records of specific religious rites which the viceroy of El-Kab was to have performed in the temples in Nubia for the king. He also appointed a man, Turi, to the position of viceroy of Cush, also known as the "King's Son of Cush." With a civilian representative of the king living in Nubia itself, Nubia did not dare to revolt as often as it had. It was easily controlled by future Egyptian kings.

Building projects

Thutmose I organized great building projects during his reign, including many temples and tombs. His greatest projects were at the Temple of Karnak under the supervision of the architect Ineni. Before this, Karnak probably consisted only of a long road to a central platform, with a number of shrines for the solar boats along the side of the road. Thutmose had the temple made much bigger. He had a fifth pylon built along the temple's main road. He built a wall around the inner temple and two flagpoles on either side of the gateway. Outside of this, he built a fourth pylon and another enclosing wall. He built a hypostyle hall between pylons four and five. This was made with columns of cedar wood. This was common in ancient Egyptian temples, and supposedly represents a papyrus marsh, an Egyptian symbol of creation. Along the edge of this room he built very large statues, each one alternating wearing the crown of Upper Egypt and the crown of Lower Egypt. Finally, outside of the fourth pylon, he built four more flagpoles and two obelisks. One of them, which now has fallen, was not inscribed until Thutmose III about 50 years later. The cedar columns in Thutmose I's hypostyle hall were replaced with stone columns by Thutmose III, however at least the northernmost two were replaced by Thutmose I himself. Hatshepsut also erected two of her own obelisks inside of Thutmose I's hypostyle hall.

Thutmose I also built statues of the Ennead at Abydos, buildings at Armant, Ombos, el-Hiba, Memphis, and Edfu, as well as minor expansions to buildings in Nubia, at Semna, Buhen, Aniba, and Quban.

Death and burial
Thutmose I was the first king who definitely was buried in the Valley of the Kings. The architect, Ineni, built the tomb, and his mortuary temple. This temple has not been found as it was probably destroyed by the building of Hatshepsut's mortuary temple at Deir el-Bahri.

Thutmose I was buried in a tomb now called KV20 dug high into the cliffs of the Valley of the Kings. His daughter Hatshepsut had the tomb made bigger, with a room big enough for both of their stone coffins (sarcophagus). His grandson, Thutmose III had him reburied in KV38. KV20 was rediscovered during the Napoleonic expedition of 1799. In 1844, the Prussian scholar Karl Richard Lepsius had explored its upper passage. However, many passageways were blocked by rocks, stones and other rubbish brought in during floods. In 1903-1904 Howard Carter, after two years of hard work, was able to clear the passages and enter the double burial chamber. He found broken pottery and shattered stone vessels from the burial chamber and lower passages. He found parts of two vases made for Queen Ahmose Nefertari, which were part of the original funerary equipment of Thutmose I. On one vase was written that Thutmose II "[made it] as his monument to his father." Other vessels which had the names and titles of Thutmose I had also been inscribed by his son and successor, Thutmose II. There were parts of stone vessels made for Hatshepsut before she herself became king which had her royal name of 'Maatkare'.

Carter, however, also discovered two separate coffins in the burial chamber. The beautifully carved sarcophagus of Hatshepsut "was discovered open with no sign of a body, and with the lid lying discarded on the floor." It is now in the Cairo Museum with a matching yellow quartzite canopic chest. A second sarcophagus, was found lying on its side, with its almost undamaged lid propped against the wall. This was given to Theodore M. Davis, who had paid for the excavation. Davis gave it to the Museum of Fine Arts in Boston. The sarcophagus had originally been engraved with the name of "the King of Upper and Lower Egypt, Maatkare Hatshepsut." However, when the sarcophagus was complete, Hatshepsut decided to have a new one made for herself. She used the first sarcophagus for her father, Thutmose I. The stonemasons then tried to erase the original carvings by restoring the surface of the quartzite so that it could be re-carved with the name and titles of Thuthmose I. The sarcophagus is seven feet long and three feet wide with walls five inches thick. It has text carved on it which records Hatshepsut's generosity towards her father:

After Hatshepsut's death, Thutmose III, Hatshepsut's successor, moved his grandfather to a magnificent new tomb, KV38. This had another yellow sarcophagus dedicated to Thutmose I. It was covered with texts which spoke of this pharaoh's love for his dead grandfather. Thutmose I's remains were disturbed during the 20th dynasty when KV38 was robbed. The sarcophagus' lid was broken and all this king's valuable precious jewellery and grave goods were stolen. The original coffin of Thutmose I was taken over and re-used by a later pharaoh of the 21st dynasty. In 1994 a flood refilled KV20 with stones and debris.

Thutmose I's mummy has not yet been identified. He was reburied during 21st dynasty when the bodies of the pharaohs of the 18th and 19th dynasties were rewrapped and buried in the Deir el-Bahri Cache above the Mortuary Temple of Hatshepsut. They were discovered in 1881. The mummies included Ahmose I, Amenhotep I, Thutmose II, Thutmose III, Ramesses I, Seti I, Ramesses II, and Ramesses IX, as well as the 21st dynasty pharaohs Pinedjem I, Pinedjem II, and Siamun. Egyptologist Gaston Maspero, believed that unlabelled mummy #5283 was that of Thutmose I. The face had the same features as the mummies of Thutmose II and Thutmose III. Later research proved that it was an 18th dynasty mummy.

Gaston Maspero wrote:

This mummy can be viewed in the Egyptian Museum in Cairo. However, in 2007, Dr. Zahi Hawass announced that the mummy is that of a 30-year-old man who had died as a result of an arrow wound to the chest. Because of the young age of the mummy and the cause of death, it could not be King Thutmose I.

References

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2nd-millennium BC pharaohs
16th-century BC births
1493 BC deaths

Thutmose I


The pars reticulata (SNpr) is a portion of the substantia nigra and is located lateral to the pars compacta. Most of the neurons that project out of the pars reticulata are inhibitory GABAergic neurons (i.e., these neurons release GABA, which is an inhibitory neurotransmitter).

Anatomy
Neurons in the pars reticulata are much less densely packed than those in the pars compacta (they were sometimes named pars diffusa). They are smaller and thinner than the dopaminergic neurons and conversely identical and morphologically similar to the pallidal neurons (see primate basal ganglia). Their dendrites as well as the pallidal are  preferentially perpendicular to the striatal afferents. The massive striatal afferents correspond to the medial end of the nigrostriatal bundle. Nigral neurons have the same peculiar synaptology with the striatal axonal endings. They make connections with the dopamine neurons of the pars compacta whose long dendrites plunge deeply in the pars reticulata. The neurons of the pars reticulata produce the neurotransmitter gamma-aminobutyric acid (GABA).
The neurons of the pars reticulata through the nigrothalamic bundle send axons to a particular part of the motor thalamus. The nigral territory corresponds to the nucleus ventralis anterior (VA) (see also List of thalamic nuclei) (different from the pallidal VO). VA is the origin of one output of the basal ganglia system. It sends axons to the frontal and oculomotor cortex. In addition the pars reticulata sends neurons to the pars parafascicularis of the central region of the thalamus and to the  pedunculopontine complex).
The particularity of the pars lateralis is to send its axons to the superior colliculus, which is a too minimized output of the basal ganglia system.

Function
The neurons of the pars reticulata are fast-spiking pacemakers, generating action potentials in the absence of synaptic input. In primates they discharge at a mean rate of 68 Hz in contrast to dopaminergic neurons (below 8 Hz). They receive abundant afferrences from the striatum (mainly from the associative striatum) with the same very peculiar synaptology as the pallidum. It receives axons from the subthalamic nucleus and a dopaminergic innervation from the dopaminergic ensemble.

The pars reticulata is one of the two primary output nuclei of the basal ganglia system to the motor thalamus (the other output is the internal segment of the globus pallidus). The nigral neurons have their own territory distinct from the cerebellar and the pallidal in the nucleus ventralis anterior VA. This sends axons to the frontal and oculomotor cortex. Hikosaka and Wurtz devoted four papers to "the visual and oculomotor functions of the monkey substantia nigra pars reticulata". This is largely involved in orientation and the control of eye movements in stabilisation of gaze and in saccades. Pars reticulata sends inhibitory projections to the superior colliculus, inhibiting eye movement, and this inhibition is 'lifted', i.e. the projecting neurons cease firing, during saccades.

Pathology
The function of the neurons of the pars compacta (not reticulata) is profoundly changed (60% of Dopamine secreting neurons, 80% decrease in dopamine in striatum) in parkinsonism and epilepsy. These changes are thought to be mostly secondary to pathology elsewhere in the brain, but may be crucial to understanding the generation of the symptoms of these disorders.

References

Basal ganglia
Midbrain

Pars reticulata