Stephen's contested accession initiated the widespread civil unrest later called the Anarchy. Count Geoffrey had little interest in England. Instead he commenced a ten-year war for the duchy of Normandy, but it became clear that to bring this conflict to a successful conclusion Stephen would need to be challenged in England so in Matilda and her half-brother, Robert invaded England. In Stephen was captured at the Battle of Lincoln and later exchanged for Robert who had also been captured.
Geoffrey continued the conquest of Normandy and in transferred the duchy to Henry while retaining the primary role in the duchy's government. Of Henry's siblings, William and Geoffrey died unmarried and childless, but the tempestuous marriage of Henry and Eleanor, who already had two daughters Marie and Alix through her first marriage to King Louis, produced eight children in thirteen years: . Henry also had illegitimate children with several mistresses, possibly as many as twelve.
Henry reasserted and extended previous suzerainties to secure possession of his inherited realm. Becket's defiance as Archbishop alienated the king and his counsellors. Henry and Becket had repeated disputes over issues such as church tenures, the marriage of Henry's brother, and taxation.
Henry reacted by getting Becket and other English bishops to recognise sixteen ancient customs in writing for the first time in the Constitutions of Clarendon , governing relations between the king, his courts and the church. When Becket tried to leave the country without permission, Henry tried to ruin him by filing legal cases relating to Becket's previous tenure as chancellor. Becket fled and remained in exile for five years.
Relations later improved, and Becket returned, but they declined again when Henry's son was crowned as coregent by the Archbishop of York , which Becket perceived as a challenge to his authority. Becket later excommunicated those who had offended him. When he received this news, Henry said: "What miserable drones and traitors have I nurtured and promoted in my household who let their lord be treated with such shameful contempt by a low-born clerk.
Henry was widely considered complicit in Becket's death throughout Christian Europe. This made Henry a pariah; in penance, he walked barefoot into Canterbury Cathedral, where he was severely whipped by monks. From Henry claimed that Pope Adrian IV had given him authorisation to reform the Irish church by assuming control of Ireland, but Professor Anne Duggan's research indicates that the Laudabiliter is a falsification of an existing letter and that was not in fact Adrian's intention.
Henry did not personally act on this until by which time William was already dead. He invaded Ireland to assert his authority over knights who had accrued autonomous power after they recruited soldiers in England and Wales and colonised Ireland with his permission. Henry later gave Ireland to his youngest son, John. This angered Henry's eighteen-year-old son, Henry the Young King , who believed these were his.
A rebellion by Henry II's wife and three eldest sons ensued. Louis VII of France supported the rebellion. William the Lion , king of the Scots, and others joined the revolt. After eighteen months, Henry subdued the rebels. In Le Mans in , Henry II gathered his children to plan a partible inheritance : his eldest surviving son, Henry, would inherit England, Normandy and Anjou; Richard his mother's favourite would inherit the Duchy of Aquitaine; Geoffrey would inherit Brittany; and John would inherit Ireland.
This resulted in further conflict. The younger Henry rebelled again, but died of dysentery. Geoffrey died in after an accident in a tournament. Henry was forced to accept humiliating peace terms, including naming Richard his sole heir. The old King died two days later, defeated and miserable. French and English contemporary moralists viewed this fate as retribution for the murder of Becket;  even his favourite legitimate son, John, had rebelled although the constantly loyal illegitimate son Geoffrey remained with Henry until the end.
Following Richard's coronation he quickly put the kingdom's affairs in order and departed on a Crusade for the Middle East. Opinion of Richard has fluctuated. Although he was respected for his military leadership and courtly manners, he had rejected and humiliated the sister of the king of France, deposed the king of Cyprus and later sold the island, he made enemies on the Third Crusade such as Leopold V, Duke of Austria by showing disrespect to his banners as well as refusing to share the spoils of war, and was rumoured to have arranged the assassination of Conrad of Montferrat.
His ruthlessness was demonstrated by his massacre of 2, prisoners in Acre. According to Steven Runciman Richard was "a bad son, a bad husband and a bad king". Returning from the crusade with a small band of followers, Richard was captured by Leopold and was passed to Emperor Henry VI. Henry held Richard captive for eighteen months — while his mother raised the ransom, valued at , marks.
After returning to England, Richard forgave John and re-established his authority in England. He left again in and battled Philip for five years, attempting to regain the lands seized during his captivity. When close to complete victory, he was injured by an arrow during a siege and died ten days later. Richard's failure to provide an heir caused a succession crisis and conflict between supporters of the claim of his nephew, Arthur , and John.
John won a significant victory while preventing Arthur's forces from capturing his mother, seizing the entire rebel leadership at the Battle of Mirebeau and his sister Eleanor, Fair Maid of Brittany. Instead he kept his prisoners so vilely and in such evil distress that it seemed shameful and ugly to all those who were with him and who saw this cruelty' according to the L'Histoire de Guillaume le Marechal.
His son, King Henry III, maintained the claim to the Angevin territories until December when he formally surrendered them and in return was granted Gascony as duke of Aquitaine and a vassal of the king of France. Sir James Holt suggests this was the inevitable result of superior French resources. John Gillingham identifies diplomatic and military mismanagement and points out that Richard managed to hold the Angevin territory with comparable finances.
By John had re-established his authority in England and planned what Gillingham has called a grand strategy to recapture Normandy and Anjou. He also brought his niece Eleanor of Brittany, aiming to establish her as Duchess of Brittany. The plan failed when John's allies were defeated at the Battle of Bouvines. Otto retreated and was soon overthrown, William was captured by the French and John agreed to a five-year truce. All subsequent English monarchs were descendants of the Angevin line via John, who had five legitimate children with Isabella : . John also had illegitimate children with several mistresses.
Thereafter, support for Louis declined, and he renounced his claims in the Treaty of Lambeth after Marshal's victories at the battles of Lincoln and Sandwich in They both overran much of Henry's remaining continental lands, further eroding the Angevins' power on the continent. In his political struggles, Henry perceived many similarities between himself and England's patron saint, Edward the Confessor. During an assembly feudal prerogatives of the king were challenged by the barons, bishops and magnates who demanded that the king reissue the Magna Carta and the Charter of the Forest in exchange for support.
Henry declared that the charters were issued of his own "spontaneous and free will" and confirmed them with the royal seal, giving the new Great Charter and the Charter of the Forest of much more authority than any previous versions. Henry III had nine children: . Henry was bankrupted by his military expenditure and general extravagance.
The pope offered Henry's brother Richard the Kingdom of Sicily , but the military cost of displacing the incumbent Emperor Frederick was prohibitive. Matthew Paris wrote that Richard stated: "You might as well say, 'I make you a present of the moon — step up to the sky and take it down'. The barons led by Henry's brother-in-law Simon de Montfort forced him to agree to the Provisions of Oxford , under which his debts were paid in exchange for substantial reforms. In France, with the Treaty of Paris , Henry formally surrendered the territory of his Angevin ancestors to Louis IX of France , receiving in return the title duke of Aquitaine and the territory of Gascony as a vassal of the French king.
Disagreements between the barons and the king intensified. De Montfort assembled the Great Parliament , recognized as the first Parliament because it was the first time the cities and boroughs had sent representatives. Louis died before Edward's arrival, but Edward decided to continue. The result was disappointing; Edward's small force only enabled him to capture Acre and launch a handful of raids.
After surviving an assassination attempt, Edward left for Sicily later in the year, never to participate in a crusade again. When Henry III died, Edward acceded to the throne; the barons swore allegiance to him even though he did not return for two years. Edward and Eleanor had sixteen children; five daughters survived to adulthood, but only one son survived Edward: .
Edward and Margaret had two sons, who both lived to adulthood, and a daughter who died as a child: . Evidence for Edward's involvement in legal reform is hard to find but his reign saw a major programme of legal change. Much of the drive and determination is likely to have come from the king and his experience of the baronial reform movement of the late s and early s.
With the Statutes of Mortmain , Edward imposed his authority over the Church; the statutes prohibited land donation to the Church, asserted the rights of the Crown at the expense of traditional feudal privileges, promoted the uniform administration of justice, raised income and codified the legal system. His military campaigns left him in heavy debt and when Philip IV of France confiscated the Duchy of Gascony in , Edward needed funds to wage war in France. When Edward summoned a precedent-setting assembly in order to raise more taxes for military finance, he included lesser landowners and merchants.
The resulting parliament included barons, clergy, knights, and burgesses for the first time. On his accession, Edward I sought to organise his realm, enforcing his claims to primacy in the British Isles. Llywelyn ap Gruffudd claimed to rule North Wales "entirely separate from" England but Edward viewed him to be "a rebel and disturber of the peace". Edward's determination, military experience and skilful naval manoeuvres ended what was to him rebellion.
The invasion was executed by one of the largest armies ever assembled by an English king, comprising Anglo-Norman cavalry and Welsh archers and laying the foundation for future victories in France. Llywelyn was driven into the mountains, later dying in battle.
Edward spent vast sums on his two Welsh campaigns with a large portion of it spent on a network of castles. Edward asserted that the king of Scotland owed him feudal allegiance, and intended to unite the two nations by marrying his son Edward to Margaret , the sole heir of King Alexander III. By invitation of Scottish magnates, Edward I resolved the dispute, ruling in favour of John Balliol , who duly swore loyalty to him and became king. Edward insisted that he was Scotland's sovereign and possessed the right to hear appeals against Balliol's judgements, undermining Balliol's authority.
Balliol allied with France in ; Edward invaded Scotland the following year, deposing and exiling Balliol. Edward was less successful in Gascony, which was overrun by the French. With his resources depleting, Edward was forced to reconfirm the Charters, including Magna Carta, to obtain the necessary funds. Meanwhile, William Wallace rose in Balliol's name and recovered most of Scotland. Wallace was defeated at the Battle of Falkirk , after which Robert the Bruce rebelled and was crowned king of Scotland.
Edward died while travelling to Scotland for another campaign. King Edward II's coronation oath on his succession in was the first to reflect the king's responsibility to maintain the laws that the community "shall have chosen" aura eslu in French. Edward's humiliating defeat by Bruce at the Battle of Bannockburn , confirming Bruce's position as an independent king of Scots, leading to Lancaster being appointed head of the king's council. The French monarchy asserted its rights to encroach on Edward's legal rights in Gascony.
Charles's sister, Queen Isabella , was sent to negotiate and agreed a treaty that required Edward to pay homage in France to Charles. Edward resigned Aquitaine and Ponthieu to his son Edward , who travelled to France to give homage in his stead. With the English heir in her power, Isabella refused to return to England unless Edward II dismissed his favourites, and she became the mistress of Roger Mortimer. Although there is no historical record of the cause of death, he is popularly believed to have been murdered at Berkeley Castle by having a red-hot poker thrust into his bowels. Mortimer was executed.
Though removed from power, Isabella was treated well, and lived in luxury for the next 27 years. In Charles IV of France died without a male heir. Queen Isabella made a claim to throne of France on behalf of her son Edward on the grounds that he was a matrilineal grandson of Philip IV of France. Not yet in power, Edward paid homage to Phillip as Duke of Aquitaine.
In Phillip confiscated Aquitaine and Ponthieu from Edward alleging he was harbouring Phillip's fugitive cousin and enemy, Robert of Artois. His army was caught by a much larger French force at Poitiers , but the ensuing battle was a decisive English victory resulting in the capture of John II of France. To reach agreement, clauses were removed that would have had Edward renounce his claim to the French crown in return for territory in Aquitaine and the town of Calais.
These were entered in another agreement to be effected only after the transfer of territory by November but both sides prevaricated over their commitments for the following nine years. Hostages from the Valois family were held in London while John returned to France to raise his ransom. When the hostages escaped back to France, John was horrified that his word had been broken and returned to England, where he eventually died. Edward and Peter fell out when Peter was unable to reimburse Edward's military expenses leaving him bankrupt.
He invaded with an army of men. Fighting was inconclusive before Gaunt agreed a treaty with King Juan of Castile. Edward's long reign had forged a new national identity, reinforced by Middle English beginning to establish itself as the spoken and written language of government. As a result, he is considered by many historians in cultural respects the first 'English' post-conquest ruler.
The Black Prince's ten-year-old son succeeded as Richard II of England on the death of his grandfather, nominally exercising all the powers of kingship supported by various councils. His government levied poll taxes to finance military campaigns and combined with the poor state of the economy resulted in the Peasants' Revolt in , followed by brutal reprisals against the rebels. The king's uncle Thomas of Woodstock, 1st Duke of Gloucester , Richard FitzAlan, 11th Earl of Arundel , and Thomas de Beauchamp, 12th Earl of Warwick , became known as the Lords Appellant when they sought to impeach five of the king's favourites and restrain what was increasingly seen as tyrannical and capricious rule.
Initially, they were successful in establishing a commission to govern England for one year, but they were forced to rebel against Richard, defeating an army under Robert de Vere, Earl of Oxford , at the skirmish of Radcot Bridge. Richard was reduced to a figurehead with little power. As a result of the Merciless Parliament , de Vere and Michael de la Pole, 1st Earl of Suffolk , who had fled abroad, were sentenced to death in their absence.
Alexander Neville, Archbishop of York , had all his possessions confiscated. Several of Richard's council were executed. On John of Gaunt's return from Spain, Richard was able to re-establish his power, having Gloucester murdered in captivity in Calais. Warwick was stripped of his title. Bolingbroke and Mowbray were exiled. When John of Gaunt died in , Richard disinherited John's son, Henry, who invaded England in response with a small force that quickly grew in numbers. Richard died in captivity early the next year, probably murdered, bringing an end to the main Plantagenet line.
None of Henry's heirs were free from challenge on the grounds of not being the true heir of Richard II and that the Lancastrian dynasty had gained the throne by an act of usurpation. Henry went to convoluted legal means to justify his succession. Many Lancastrians asserted that his mother had had legitimate rights through her descent from Edmund Crouchback , who it was claimed was the elder son of Henry III of England, set aside due to deformity.
The later marriage of his sister Anne to Richard of Conisburgh, 3rd Earl of Cambridge consolidated this claim to the throne with that of the more junior House of York. Henry planned to resume war with France, but was plagued with financial problems, declining health and frequent rebellions. Henry IV died in His son and successor, Henry V of England , aware that Charles VI of France 's mental illness had caused instability in France, invaded to assert the Plantagenet claims and won a near total victory over the French at the Battle of Agincourt. The resulting Treaty of Troyes stated that Henry's heirs would inherit the throne of France, but conflict continued with the Dauphin.
Humphrey's wife was accused of treasonable necromancy after two astrologers in her employ unwisely, if honestly, predicted a serious illness would endanger Henry VI's life, and Humphrey was later arrested and died in prison. Depopulation stemming from the Black Death led to increased wages, static food costs and a resulting improvement in the standard of living for the peasantry. However, under Henry misgovernment and harvest failures depressed the English economy to a pitiful state known as the Great Slump.
The economy was in ruins by , a consequence of the loss of France, piracy in the channel and poor trading relations with the Hanseatic League. Both of Edmund's sons were killed in Richard became involved in the Southampton Plot , a conspiracy to depose Henry V in favour of Richard's brother-in-law Edmund Mortimer. When Mortimer revealed the plot to the king, Richard was executed for treason. Richard's childless older brother Edward was killed at the Battle of Agincourt later the same year.
The increasingly interwoven Plantagenet relationships were demonstrated by Edmund's second marriage to Joan Holland. Margaret Holland , another of Joan's sisters, married John of Gaunt's son. Anne died giving birth to their only son in September Although his earldom was forfeited, Richard the father was not attainted , and the four-year-old orphan Richard was his heir. Within months of his father's death, Richard's childless uncle, Edward Duke of York, was killed at Agincourt.
Richard was allowed to inherit the title of Duke of York in In he acquired the earldoms of March and Ulster on the death of his maternal uncle Edmund Mortimer, Earl of March, who had died campaigning with Henry V in France, and the earldom of Cambridge which had belonged to his father. Being descended from Edward III in both the maternal and the paternal line gave Richard a significant claim to the throne if the Lancastrian line should fail, and by cognatic primogeniture arguably a superior claim.
Having inherited the March and Ulster titles, he became the wealthiest and most powerful noble in England, second only to the king himself. Richard married Cecily Neville , a granddaughter of John of Gaunt, and had thirteen or possibly fifteen children: . When Henry VI had a mental breakdown, Richard was named regent, but the birth of a male heir resolved the question of succession.
The ruling class was deeply shocked and reconciliation was attempted. Margaret found this disregard for her son's claims unacceptable, and so the conflict continued. York was killed at the Battle of Wakefield and his head set on display at Micklegate Bar along with those of Edmund, Earl of Rutland , and Richard Neville, Earl of Salisbury, who had been captured and beheaded.
He was crowned after consolidating his position with victory at the Battle of Towton. Edward's preferment of the former Lancastrian-supporting Woodville family, following his marriage to Elizabeth Woodville , led Warwick and Clarence to help Margaret depose Edward and return Henry to the throne. Edward and Richard, Duke of Gloucester , fled, but on their return, Clarence switched sides at the Battle of Barnet , leading to the death of the Neville brothers.
The subsequent Battle of Tewkesbury brought the demise of the last of the male line of the Beauforts. By the mids, the victorious House of York looked safely established, with seven living male princes: Edward IV and his two sons, his brother George and George's son, his brother Richard and Richard's son. Edward and Elizabeth Woodville themselves had ten children, seven of whom survived him: . Dynastic infighting and misfortune quickly brought about the demise of the House of York. George Plantagenet, 1st Duke of Clarence , plotted against his brother and was executed.
Following Edward's premature death in , his brother Richard had Parliament declare Edward's two sons illegitimate on the pretext of an alleged prior pre-contract to Lady Eleanor Talbot , leaving Edward's marriage invalid. Richard's son predeceased him and Richard was killed in  after an invasion of foreign mercenaries led by Henry Tudor , who claimed the throne through his mother Margaret Beaufort. When Henry Tudor seized the throne there were eighteen Plantagenet descendants who might today be thought to have a stronger hereditary claim, and by this number had been increased further by the birth of sixteen Yorkist children.
She was the eldest daughter of Edward IV, and all their children were his cognatic heirs. Indeed, Polydore Vergil noted Henry VIII's pronounced resemblance to his grandfather Edward: "For just as Edward was the most warmly thought of by the English people amongst all English kings, so this successor of his, Henry, was very much like him in general appearance, in greatness of mind and generosity and for that reason was the most acclaimed and approved of all.
This did not deter Margaret of York, Duchess of Burgundy —Edward's sister and Elizabeth's aunt—and members of the de la Pole family—children of Edward's sister and John de la Pole, 2nd Duke of Suffolk — from frequent attempts to destabilise Henry's regime. John de la Pole, 1st Earl of Lincoln , joined the revolt, probably anticipating that it would further his own ambitions to the throne, but he was killed in the suppression of the uprising at the Battle of Stoke Field in Edward's execution may simply have been a precondition for the marriage of Arthur, Prince of Wales to Katherine of Aragon in John de la Pole's attainder meant that his brother Edmund inherited their father's titles, but much of the wealth of the duchy of Suffolk was forfeit.
Edmund did not possess sufficient finances to maintain his status as a duke, so as a compromise he accepted the title of earl of Suffolk. Financial difficulties led to frequent legal conflicts and Edmund's indictment for murder in He fled with his brother Richard, while their remaining brother, William, was imprisoned in the Tower—where he would remain until his death 37 years later—as part of a general suppression of Edmund's associates Philip the Fair had been holding Edmund and in he returned him to Henry.
Edmund was imprisoned in the Tower. In , he was executed after Richard de la Pole, whom Louis XII of France had recognised as king of England the previous year, claimed the kingship in his own right. By then, the cause was more religious and political rather than dynastic.
The attainder of her father, Clarence , was a legal bar to any claims to the throne by his children. Nevertheless, it did allow the couple to be closely involved in court affairs. This made her the first and, apart from Anne Boleyn , the only woman in 16th-century England to hold a peerage title in her own right. Buckingham's fall after arguments with the king over property, and Margaret's open support for Catherine of Aragon and Princess Mary began the Poles' estrangement from the king.
In evidence came to light that Pole family members in England had been in communication with Reginald. Margaret's sons Geoffrey and Henry were arrested for treason along with several friends and associates, including Henry's wife and brother-in-law— Edward Neville. Among those arrested was the king's cousin Henry Courtenay, 1st Marquess of Exeter , his wife and year-old son. Courteney's wife was released two years later, but their son spent 15 years in the Tower until Queen Mary released him.
Except for the surviving Geoffrey Pole, all the others implicated were beheaded. Margaret was attainted. The possibility of an invasion involving Reginald via her south coast estates and her embittered relationship with Henry VIII precluded any chance of pardon. However, the decision to execute her seems a spontaneous, rather than a premeditated, act. According to the Calendar of State Papers her execution was botched at the hands of "a wretched and blundering youth His father failed in his rebellion against Richard III in but was restored to his inheritance on the reversal of his father's attainder late in There is no evidence of continuous hostility between Buckingham and Henry VIII, but there is little doubt of the duke's dislike of Thomas Wolsey , whom he believed to be plotting to ruin the old nobility.
Buckingham himself was arrested in April ; he was found guilty on 16 May and executed the next day. Evidence was provided that the duke had been listening to prophecies that he would be king and that the Tudor family lay under God's curse for the execution of Warwick. Much of this evidence consisted of ill-judged comments, speculation and bad temper, but it underlined the threat presented by Buckingham's descent. As late as , with the Tudor succession in doubt, older Plantagenet lines remained as possible claimants to a disputed throne, and religious and dynastic factors gave rise to complications.
Thomas Wilson wrote in his report The State of England, Anno Domini that there were 12 "competitors" for the succession. At the time of writing about , Wilson had been working on intelligence matters for Lord Buckhurst and Sir Robert Cecil. Ranulph Crewe , Chief Justice of the King's Bench , argued that by the House of Plantagenet could not be considered to remain in existence in a speech during the Oxford Peerage case, which was to rule on who should inherit the earldom of Oxford.
Crewe said:. I have labored to make a covenant with myself, that affection may not press upon judgment; for I suppose there is no man that hath any apprehension of gentry or nobleness, but his affection stands to the continuance of a house so illustrious, and would take hold of a twig or twine-thread to support it. And yet time hath his revolutions; there must be a period and an end to all temporal things — finis rerum — an end of names and dignities, and whatsoever is terrene; and why not of de Vere?
For where is Bohun? Where is Mowbray? Where is Mortimer? Nay, which is more, and most of all, where is Plantagenet? They are entombed in the urns and sepulchres of mortality! Angevins  Henry II of England , —, had 5 sons;. Plantagenets . House of Lancaster . House of Beaufort illegitimate branch of House of Lancaster .
House of York . From Wikipedia, the free encyclopedia. Royal dynasty in medieval England. For other uses, see Plantagenet disambiguation. Armorial of Plantagenet. Further information: Angevin kings of England and Angevin Empire. See also: Angevin kings of England. Further information: The Anarchy.
Gwynedd , Llywelyn ap Gruffudd 's principality. Territories conquered by Llywelyn. Territories of Llywelyn's vassals. Lordships of the Marcher barons. Lordships of the king of England. Main article: House of Lancaster. Further information: Hundred Years' War — Main article: House of York. Further information: Wars of the Roses. Main article: Succession to Elizabeth I of England. See also: Plantagenet monarchs' family tree. Family tree. This family tree includes selected members of the House of Plantagenet who were born legitimate. House of York.
List of members of the House of Plantagenet all male and some significant females. Angevins  Henry II of England , —, had 5 sons; 1. William IX, Count of Poitiers , —, died in infancy 2. Henry the Young King , —, died without issue 3. Richard I of England , —, died without legitimate issue 4. Geoffrey, Duke of Brittany , —, had 1 son; A. Arthur I, Duke of Brittany , —, died without issue 5. John of England , —, had 2 sons; Plantagenets  A.
Edward I of England , —, had 6 sons. John of England, —, died young b. Henry of England, —, died young c. Alphonso, Earl of Chester , —, died young d. Edward II of England , —, had 2 sons; i. Edward III of England , —, had 8 sons; 1. Edward, the Black Prince , —, had 2 sons; A. Edward, —, died young B.
Richard II of England , —, died without issue 2. William of Hatfield, —, died in infancy 3. Lionel of Antwerp, 1st Duke of Clarence , —, 1 daughter. Anne de Mortimer , —, married Richard of Conisburgh, 3rd Earl of Cambridge see below and it is through her descent from Lionel that the House of York claimed precedence over the House of Lancaster. To the House of York  b.
Thomas of England, —, died in infancy 7. William of Windsor, —, died in infancy 8. Thomas of Woodstock, 1st Duke of Gloucester , —, had 1 son; A. Humphrey Plantagenet, 2nd Earl of Buckingham, —, died without issue ii. John of Eltham, Earl of Cornwall , —, died without issue e. Thomas of Brotherton, 1st Earl of Norfolk , —, had 2 sons; i. Edward of Norfolk, —, died young ii.
John Plantagenet, —, died without issue f. Edmund of Woodstock, 1st Earl of Kent , —, had 2 sons; i. Edmund Plantagenet, 2nd Earl of Kent, —, died young ii. Edmund Crouchback, 1st Earl of Lancaster , —, had 3 sons; a. Thomas Plantagenet, 2nd Earl of Lancaster , —, died without issue b. Henry, 3rd Earl of Lancaster , —, had 1 son; i.
Henry of Grosmont, 1st Duke of Lancaster , —, died without male issue, 2 daughters Maud, Countess of Leicester , —, died without issue Blanche of Lancaster , married John of Gaunt and had 1 son and two daughters To House of Lancaster c. Richard of England, —, died young IV. John of England, —, died young V. William of England, —, died young VI. Henry of England, —, died young B. Richard, 1st Earl of Cornwall , —, had 5 sons; I.
John of Cornwall, —, died in infancy II.
Former British citizens killed by drone strikes after passports revoked
Henry of Almain , —, died without issue III. Nicholas of Cornwall, —, died in infancy IV. Richard of Cornwall, —, died in infancy V. Edmund, 2nd Earl of Cornwall , —, died without issue House of Lancaster  4. John of Gaunt, 1st Duke of Lancaster , —, had 4 sons; A. John of Lancaster, —, died in infancy B. Edward Plantagenet, —, died in infancy C. John Plantagenet, —, died in infancy D. Henry IV of England , —, had 5 sons; I. Edward Plantagenet, —, died in infancy II. Henry V of England , —, had 1 son; a.
Henry VI of England , —, had 1 son; i. Edward of Westminster , —, died without issue III. Thomas, Duke of Clarence , —, died without issue IV. The apple WGS will also assist in understanding the effects of plant growth regulators on flowering in apple, potentially leading to development of novel compounds that could increase commercial production efficiency.
For example, the phytohormone gibberellic acid GA has long been known to repress flowering in apple, in contrast to its well-known promotion of flowering in rosette herbaceous plants , It was recently reported that exogenous GA applied to apple trees early in the growing season was associated with increased expression of MdTFL , one of two copies of the TFL1 gene, late in the growing season, at a time when flowers would be forming MdTFL1 can repress flowering when expressed ectopically in Arabidopsis and apple , and, if MdTFL1 does act as a flowering repressor in apple, then its promotion by GA would provide a simple explanation for the repression of flowering by this phytohormone.
The reference WGS will also provide insights into the genetic mechanisms underlying juvenility—arguably the most significant challenge for rapid cultivar development Indexing the full complement of apple genes, together with the finding that some well-studied mechanisms of phase change in maize and Arabidopsis are conserved in woody perennial plants , provides a smooth route for developing apple breeding germplasm to enable the rapid introduction of novel phenotypes in response to rapidly changing production environments and market demands.
In addition, an improved genomic knowledge of apple will shed light onto the interesting variance in juvenile period length observed among Malus wild and cultivated accessions. The rapid-cycling, wild apple species M. However, unlike most M. The high-quality M.
The apple WGS has been a fundamental resource for the determination of key genes involved in the climacteric response and cell wall modification of apple fruit. The attractiveness of apple fruit is due to an array of features shape, size, color, texture, and aroma , which from the initial phase of fruit development continuously changes through the fruit ripening process In climacteric fruit, including apple, fruit ripening is triggered and coordinated by the action of the hormone ethylene.
Following synthesis, this hormone is perceived by a series of receptors ethylene response sensor and ethylene resistant that initiate a downstream signaling mitogen-activated protein kinase cascade constitutive triple response—ethylene insensitive 2 activating finally a series of ethylene-dependent responses mediated by ethylene response factor transcription factors , The ethylene competitor 1-methylcyclopropene 1-MCP has commonly been used to reveal the role of ethylene in controlling the overall ripening process in apple.
Through use of these tools, a gene de-repression or de novo activation, was revealed following 1-MCP treatment Gene annotation and prediction using the apple WGS enabled detection of possible cross-talk between ethylene and auxin. Upregulation of these genes was further validated through a RNA-seq survey performed to investigate the physiological mechanisms controlling apple superficial scald Gene annotation also facilitated the detection of important elements involved in postharvest ripening, as well as in the control of this scald.
This research identified a specific pectin methylesterase MdPME2 as well as elements involved in hemicellulose metabolism , The same tool enabled a GWAS that revealed the genetic relationship between texture and the production of volatile organic compounds Further improvement of the quality of the genome assembly and fine-tuning of the gene annotation should enable more precise and informative characterization of the relevant genes involved in the control of important physiological pathways underlying the apple fruit ripening processes, ultimately enabling the development of new cultivars characterized by superior fruit quality.
Fruit appearance is a key factor in consumer acceptance Red coloration in the skin and flesh of apple fruit has been demonstrated to be controlled genetically by allelic variation , , , , by the environment differences in light and temperature , as well as by management e. MYB1 and MYB10 , which are likely to be allelic , regulate the activity of the genes of the anthocyanin pathway and their expression correlates with anthocyanin accumulation. A locus closely linked to MYB10 has been mapped and associated with red flesh and red skin using biparental populations and GWAS in germplasm collections 13 , 83 , , , , Interestingly, this red fruit R f locus is orthologous with red color-imparting loci detected in other Rosaceae fruit crops such as cherry, peach and strawberry Recently, a robust allele-specific qPCR marker was developed and validated using both apple breeding families and genetically diverse cultivars However, the causative variant for red flesh coloration compared to the usual white flesh color has been identified.
The availability of the new GDDH13 reference genome 8 , together with whole genome re-sequencing data for a range of apple cultivars old and modern 10 , 57 , opens the prospect of pinpointing the causative variant s for red skin color. More re-sequencing, including using bisulfite-treated DNA, might shed new light on the key regulatory motifs. Such work will need to be complemented with a test of the efficiency of contrasting-effect haplotypes for anthocyanin gene activation, activation of MYB10 itself, and as targets of transcription factors regulating MYB One key consideration is that red fruit skin might be associated with more than one allele.
In this case, a combination of PBA using a population segregating for red coloration derived from several ancestral sources of red determinism combined with high-throughput genome re-sequencing would be required. Fruit malate content varies widely in Malus accessions: from 0.
Enzymes and their coding genes associated with malate biosynthesis and degradation were previously emphasized, until the important role was uncovered of malate transport into the vacuole and its maintenance there , , Major differences in apple fruit acidity among cultivars were determined to be controlled by a single locus that was mapped to one end of chromosome 16 and designated as the Malic acid Ma locus , , Expression of Ma1 co-segregated with malate content and at least one allele sufficed for increased Ma1 expression, causing a three-fold increase in fruit malate content Exploration of the apple WGS enabled major discoveries after more than a decade of stagnation in characterizing the actual gene underlying apple fruit acidity , , The Ma locus, explaining Finally, a G-to-A mutation was identified at bp of the open reading frame in the Ma1 gene that leads to a premature stop codon, truncating 84 amino acids of an aluminum-activated malate transporter MdALMTII protein, and significantly reducing fruit acidity when occurring as a homozygote mama.
This is not surprising, as other acidity QTLs have been reported which occasionally had a size similar to Ma1. A QTL on LG 8 exhibited a similar or higher contribution to fruit acidity in many other segregating families when compared with the LG 16 QTL 29 , , , , although also a substantial lower contribution has been reported This difference in performance might be due to different functional alleles or tightly linked QTLs of different effect segregating among families Future research targets utilizing the apple WGS include development of a more detailed network of apple fruit malate metabolism and discovery of further natural genetic variation in the network in diverse Malus germplasm resources.
Emphasis should be on the molecular regulation pathways of malate transport to the vacuole, vacuole malate sequestration and depletion during fruit development and post-harvest storage, the effect of different growing environments and functional mutations among cultivars and species. The availability of the apple WGS has revolutionized the approaches and reduced the time needed to identify candidate resistance genes. Just over 20 years ago, the mapping of a resistance gene R gene , which is the first step towards the positional cloning of a gene, was a long and tedious endeavor.
Around the beginning of the new millennium, genetically mapped SSR markers for apple became available , , , The establishment of genetic map positions of these resistance genes opened up their potential for cloning and determination of their DNA sequence. In this case, the apple WGS allowed a rapid saturation with markers of the regions containing the R gene. The time savings can also be recognized by the increase in number of published reports of such research. Finally, the WGS facilitated determination of the diversity and identity of QTLs from different studies for the same linkage group by clarifying their relative chromosomal position However, identification of many candidate R -gene allele s cannot be achieved using the currently available apple WGS, because these resources have not been developed from individuals in the germplasm that carry the resistance alleles.
Until now, R -gene alleles have been identified by sequencing BAC clones from libraries of resistance allele-carrying individuals spanning the R -gene regions e. The fast progress in sequencing technology and sequence assembly associated with decreasing sequencing cost will soon make the use of BAC libraries obsolete. Instead, whole genome resequencing of individuals carrying an R allele of interest and the subsequent assembly of the specific region carrying that R allele is expected to become the new standard. Furthermore, determining the correct order of repetitive genes will be very important, because resistance genes are often arranged in tandem repeats With the WGS of apple speeding up the process of identifying R genes, translation to crop improvement is also increasing.
Genetic markers and knowledge gained from R -gene cloning research is allowing the building of pyramids of R alleles against the same pathogen with different mechanisms of resistance. Such pyramids of R alleles are expected to lead to the development of cultivars with more durable resistances. However, for some diseases a more straightforward approach might be the targeted knock-out of susceptibility S genes, such as for MdMLO19 that should lead to development of powdery mildew resistant apple cultivars, at least in the absence of other MLO genes of similar effect Rapid advances in apple breeding were projected in the publications of both the first and second iterations of the apple WGS 1 , 8.
Indeed, progress is being made in the development and application of genomic technologies in the area of trait-predictive tests, although perhaps not as quickly as hoped for by earlier reviewers , The use of SNP arrays 10 , 11 , 12 and GBS to efficiently screen mapping populations and construct relatively dense genetic maps 13 , has made the mapping of trait loci much less challenging and correspondingly the number of QTLs in the Genome Database for Rosaceae has risen rapidly since the availability of the apple WGS.
The recent development of a tool utilizing apple resequencing data from breeding germplasm, for the purpose of identifying SNPs that are unique to the accession carrying an attribute of interest, has both increased the efficiency of genetic mapping of trait loci and provided a source of SNPs for use in designing transferable high-throughput markers in combination with accurate pedigree records or PBA 74 , Highly reliable diagnostic markers derived from candidate genes for traits can now be readily developed by examining the DNA sequence at MTLs such as major resistance loci and the sequence underlying QTLs , Early adopters of trait-predictive tests mainly employed simple sequence repeat and SCAR markers , , , , and highly informative SSR markers are still used for tracing pedigrees 67 , Currently, high-throughput SNP-based markers have become the first choice for use in trait-predictive tests for parental and seedling selection by breeders and many programs have moved to develop and introduce the use of such tests 80 , , , , , In some families, MAS is applied for additional quality traits, such as red flesh Myba.
Screens applied vary according to year and genetic background of seedlings, but include dwarfing Dw1 , Dw2 resistances to fire blight FB-R5 , European canker Rnd1 , and woolly apple aphid Er1 , Er2 , Er3 , as well as adventitious rooting MAS targeting trait loci for fruit quality and disease resistance is also in routine use in several other apple breeding programs around the world MAS can be and is also routinely employed to reduce numbers of seedlings prior to evaluation by genome-wide selection Other types of high-throughput markers have been developed for apple, e.
Various avenues of future work are warranted to facilitate the application of trait-predictive tests for MAS. Individual breeding programs will wish to develop markers for specialized traits that are relevant to their own germplasm and breeding goals. New marker options need to be investigated for apple, e. Additional development of haplotyping, already initiated with SNP arrays, could be made using re-sequencing and tools such as Beagle Apple was the first horticultural tree crop in which this approach was explored , and implemented The premise of GP is that anonymous genome-wide markers are sufficiently dense so that most or all QTLs are in LD with a marker and that most or all QTLs segregate with the use of multi-parent populations.
Consistently, GP accuracy has been demonstrated to be higher than using large-effect QTL-only models for polygenic traits reviewed by Crossa and co-workers Results from apple studies suggest important factors leading to higher accuracy are close genetic relationships between training and validation populations , , large training population sizes , , small effective size of training and validation populations , , high heritability of traits particularly obtained through objective phenotyping , , assessment of continuously distributed traits , and high density of markers However, multiple assessments across years of the same experimental unit i.
The extent of LD in a population is related to effective population size, hence LD and genetic relationship levels are entangled in a population, making it difficult to separate their effects on the accuracy of GP. Hence, the accuracy of GP over successive generations would reduce unless GP models are recalibrated, although the equivalent is also needed with any linkage-based prediction method To date, population sizes are considerably smaller, and density of genotyping lower, in apple studies compared to those for animals.
The influence of using different prediction models has been explored in apple. Assumption of a Gaussian distribution i.
Genomic prediction models can combine whole-genome prediction and detection of large-effect QTLs , , and in apple a Bayesian LASSO model was used to detect regions of high influence for fruit quality traits The Gaussian framework, however, allows extension to more complex models and has been used to model both additive and non-additive genetic effects, although negligible effect on prediction accuracy of breeding value additive or clonal values was reported in apple and cherry The fundamental aim of GP is greater gain at reduced cost and in less time than conventional strategies and considerable increases in gain across animals and crops has been demonstrated However, as many traits are of interest in apple , this strategy might work best when focusing on a single trait that is governed by many loci of minor effect.
Selection of parents from the training population provides a strong genetic link with the selection population, to improve prediction accuracy. GP is reportedly in initial use in US apple programs J. Luby, Uni. Minnesota, pers. As GP is a unifying technology across animal and plant breeding , advances across these systems can provide direction for fundamental discoveries and practical application in apple. The ability to clone apple individuals provides opportunities to counteract low prediction accuracy due to small sample sizes or large experimental variation.
Improvement in accuracy from multi-environment and multi-trait prediction models found in other crops , , needs to be verified for apple. The biallelic nature of SNPs suggests combining linked SNPs into haplotype blocks haploblocks is likely to be more informative of the variability at a QTL and could decrease hidden variability, hence leading to higher prediction accuracy , , While the use of sequence information to build GP models suggests higher accuracy because the causal variation should be directly included in the model, the value of this extension in apple needs to be verified as populations are highly structured 69 such that large haploblocks could be efficiently tagged with a lower density of markers , Fast-track breeding via transgenesis , or induced through a plant virus vector might enable full realization of GP potential in apple.
While SSRs were used to select for a favorable QTL allele and against unfavorable genetic background over five generations in seven years , GP can be used to identify elite progeny across a range of traits. Implementation of GP for apple improvement might require modification of breeding programs and stochastic simulation can be used to evaluate benefits of different strategies This approach can therefore leverage the latent value of historically collected trial data. Bioinformatics tools are being developed to manage and deliver the technology so that the performance of new germplasm can then be routinely predicted.
Access to a near-complete apple WGS provides the foundation for epigenetic studies. By definition, epigenetic changes result in heritable changes in gene expression that cannot be explained by changes in the DNA sequence. To be able to study true epigenetic events, one has to ensure that there are as few genetic changes as possible in the genome under examination.
Therefore, apple sports are therefore an excellent starting point to study epigenetics in plants. Although some sports might be the result of genetic changes such as mutations induced by transposable elements , others might vary only at the epigenetic level and more specifically at the DNA methylation level.
The main reason for generating the apple reference genome GDDH13 was to enable an understanding of the epigenetic mechanisms involved in regulating fruit size. This approach was successful, as the research generated a list of genes likely involved in fruit size regulation 8. Currently, there is no direct impact of apple epigenetics research on crop improvement. Future studies using epigenetic markers on sports and on larger populations will enable assessment of the contribution of DNA methylation to traits of interest. Considering that DNA methylation also influences fruit color intensity and patterning in apple sports , , it is likely that we have only started to scratch the surface of the contribution of epigenetics to traits of economic importance.
Several epigenetic studies are expected to profit from the availability of high-quality apple WGSs in the near future. The study of the genetic and epigenetic mechanisms influencing phenotypic changes of apple sports is straightforward with a good genome assembly and will allow the rapid identification of genes involved in important agronomic traits, including fruit color and shape, disease resistance and tree architecture.
Chemical-induced or targeted changes in DNA methylation patterns could be used to produce novel traits of interest by gene demethylation. DNA methylation changes can also be used to induce the artificial mobilization of transposable elements , which results in a powerful tool for gene discovery via tagged mutations and the generation of phenotypic diversity.
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The GDR is built using Tripal , , a resource-efficient, open-source platform for online biological database construction. Increased data from the worldwide apple research community, combined with active curation, analyses, and tool development, has resulted in significant expansion of the GDR during the last few years. Described below are the currently available data and interfaces, with a focus on new features. The physical structure of the apple WGS, as described by its nucleotide sequence, is the foundation of apple genomic resources in the GDR. Although no chromosomal sequence is available for these heterozygous apple genome assemblies, the sequences of overlapping contigs are available, which in turn have been aligned to the chromosome using markers from the genetic map.
For the Golden Delicious v1. Researchers can readily examine in fine detail how the apple genome compares physically with those of other crops. The most recent apple WGS is being used in synteny analysis with eight other Rosaceae whole genome assemblies using MCScanX , with the results available through the new Synteny Viewer. Another strong focus of the GDR is apple gene function. Since , the community database has provided an apple reference transcriptome M. Genetic linkage, marker and trait locus information needs are well served.
The GDR contains genetic maps for Malus , which can be viewed and compared through a new graphic interface, MapViewer. Currently, the GDR contains 2. Most of the remaining SNPs are those identified in silico from accession resequencing conducted during development of those SNP arrays. Although impressive, some caution is necessary in considering these numbers.
They are based on the accumulation of QTLs as independently reported in the literature and therefore might include redundancies across studies or treatments including years within a single study. In addition, some caution is be needed in using reported QTL information. Their assignment might arise from different levels of statistical significance; regrettably, it is often not specified whether a genome-wide or a chromosome-wide threshold was applied.
Some reported QTLs were from an interval mapping procedure without subsequent co-factor analysis. Such QTLs have increased risk of being spurious or being assigned to an incorrect position, while other true QTLs might remain hidden , BIMS allows breeders to store, manage, archive, and analyze their private or public breeding data.
Functional annotation of gene models using sequence similarity and synteny analyses provided a first step in data integration across species. Alignment of transcripts and markers, used in genotyping and QTL mapping, to the WGS, enables researchers to utilize data of different types originating from various species and disciplines. Future targets include integrating new types of data such as pan-genome data, epigenome data, and phenomics data, as well as data standardization and further tool development.
In this way, the GDR will continue to enable researchers to open new frontiers in understanding the structure and function of the apple genome and its components. Over the past decade, apple WGSs have had an enormous impact on our understanding of apple biological functioning, trait physiology and inheritance, leading to practical applications for improving this highly valued crop.
The first apple WGS reported in 1 , v1. As the research community exploited this resource, the need for improvements in the genome assembly became increasingly apparent. The new GDDH13 assembly reported in 8 provided many of those improvements, and research leaps ensued.
In contrast to a decade ago, causal gene identities for phenotypes of fundamental or practical interest can today be discovered rapidly. Genome-wide polymorphisms are readily screened for hundreds to thousands of germplasm individuals. Availability of the WGS is not only turbo-charging the conduct of these classical research steps to crop improvement, but is also driving innovative methods of achieving more durable, environmentally sound, productive, and consumer-desirable apple crop production. Research collaboration on apple genomics and genetics, already vibrant across an international community, has much synergistic value.
Although it can be viewed in many ways, each chromosome has the same physical entity for any researcher. Researchers worldwide are resolutely identifying the roles and influences, intra-genomic interactions, and germplasm distributions of variants of chromosomal modules—the genes, motifs, trait loci, haploblocks, base pairs and so on—all using the WGS of apple. Integration of the apple transcriptome, proteome, metabolome, epigenome, QTLome , and so on, across experiments and experimental material will surely expand our scientific understanding of apple.
The sheer amount of multi-dimensional genomic and downstream data generated has elevated the need and opportunities for their coordination to a new level. To sustain such collaborative advances, the GDR now serves as the research hub for the international apple genomics community. We should also think beyond apple: exploiting the genomic synteny of apple with crop relatives in the Rosaceae family has only just begun, and holds much potential for wider collaboration and research breakthroughs. Advances are needed in upgrades to the base genome sequence data pool. The reference WGS should be improved to ensure its completeness and accuracy.
A first step could be noting in genome browsers any suspicious regions identified by comparisons between the WGS and high-quality genetic linkage maps. Given the possibility that the doubled-haploid individual underlying the current reference WGS is missing parts of its genome compared to typical heterozygous cultivars 8 , there is still a need for a reference WGS representing the complete cultivated apple genome.
Development of the apple pan-genome would be a clear way forward for describing the current allelic variation of cultivated apple. Expansion of the pan-genome to encompass the Malus genus is a further compelling target as M. Advances are also needed in use of WGS-related data to answer fundamental and applied research questions. Genomic prediction approaches promise a foundation for genotype and environment matching. A major leap in analytical capabilities is required to gain information on the vast reservoir of alleles that already contribute to the phenotypes of cultivated and wild apple trees.
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