Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

On November 7, 1491, the Peace of Presburg was signed, which ended the decades-long war between Hungary and Austria [14].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

A new stage in the expansion of the Austrian state is also associated with the name of Maximilian I: in 1515 he organized a meeting in Vienna with Vladislav II, King of Hungary and the Czech Republic, and Sigismund I, King of Poland, representatives of the Jagiellon dynasty.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

An agreement was concluded on the marriage of the granddaughter of Maximilian I Maria and the son of Vladislav II Lajos II, as well as on the marriage of the grandson of Maximilian Ferdinand to the daughter of Vladislav II Anna.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

This dynastic union made it possible in 1526, after the death of Louis II in the war with the Turks, to annex Hungary and the Czech Republic to the possessions of the Habsburg [15].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Bavarian Succession Disputes


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Georg the Rich, Duke of Lansgut-Bavaria, died in 1503.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Not wanting his possessions to go to Albrecht IV of Bavarian-Munich, he made a will shortly before his death, in which Ruprecht of the Palatinate, married to his daughter Elizabeth, made the heir.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

However, Maximilian at one time signed an agreement with Albrecht, who was guaranteed the inheritance of Lansgut.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

As a result, Albrecht started a war against Ruprecht, who occupied Lansgut on behalf of his wife.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Albrecht justified his rights by the fact that in Germany, inheritance of possessions through the female line violates existing laws.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

All Maximilian's attempts to resolve the conflict peacefully failed [16].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In the outbreak of the war for the Landsgut inheritance, Maximilian sided with Albrecht, the husband of his sister.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

He outlawed Ruprecht, which only fanned the conflict.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Ruprecht was supported by his father, Elector Philip of the Palatinate, as well as the King of Bohemia.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The war went on with varying success, devastating the affected territories.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In August 1504, Ruprecht died at the Battle of Lansgut.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

However, peace came only in September after in the Battle of Wenzenbach (near Regensburg), the army led by Maximilian defeated the Palatinate and Czech troops, and Maximilian, who miraculously escaped death, proved to be a brave warrior [16].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The result of the war was that Albrecht IV united the Bavarian lands in his hands.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

At the same time, Maximilian annexed to his possessions a part of Tyrol that previously belonged to Bavaria (Kufstein, Rattenberg, Kitzbühel, Zillertal, as well as the area around Mondsee).


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

This completed the folding of the core of the Austrian state of modern times.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Once again, Maximilian had to intervene in Bavarian affairs after the death of Albrecht IV in 1508.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Shortly before his death, Albrecht issued a new law on the inheritance of Bavarian lands by birthright.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Fearing conflicts between her sons, Albrecht Kunigund's widow asked her brother to change the law by allowing inheritance to the youngest sons, which Maximilian did.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

However, as a result, both youngest sons did not leave children [16].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian I's Reforms


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian's reign was of great importance for the development of the Austrian state.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The Archduke launched a broad program for transforming the country's public administration system.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Already in 1493, Austrian lands were divided into two districts: Upper Austria (Tyrol and Western Austria) and Lower Austria (Austria, Styria, Carinthia, Kraina).


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In each of the parts, a separate government was created, including a governor appointed by the archduke and his advisers.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Vienna organized a single treasury for all Länder (later moved to Innsbruck) and an accounts office.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In 1498 Maximilian created a whole system of supreme governing bodies: the Court Council (the highest administrative and judicial body), the Court Chamber (the highest fiscal and financial body) and the Court Chancellery ( external relations and legislative activity).


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The military departments of all Austrian states were also united.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

As a result, a new, centralized system of public administration was organized and the basis for the unification of the ancestral Habsburg possessions into a single Austrian state was laid.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The reforms of Maximilian I, however, caused resistance from the estates.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In 1502, in a single Diet of the Lower Austrian lands, the nobility strongly opposed the centralization aspirations of the archduke.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian, who is in dire need of funds to wage wars with the Turks and in Italy, was forced to make concessions and restore the judicial rights of the estates.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In 1518, representatives of the Diet were included in the Court Council.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Thus, administrative reforms were not completed, but they contributed to the strengthening of state power in Austria and made it possible to create an effective bureaucratic system in the future.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The rule of the Holy Roman Empire.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Imperial reform


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Unlike his father, the indecisive and unlucky Frederick III, Maximilian already in his young years proved to be an energetic statesman capable of strengthening royal power and exercising urgent transformations of the archaic state systems of Germany and Austria.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Already in the 1480s, Emperor Frederick III, who reached the age of 70, actually withdrew from the management of his possessions, handing over the reins of power to his son.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

And in 1486, Friedrich gathered six electors in Frankfurt (only the king of Bohemia was absent), who elected Maximilian King of Germany on February 16.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

At the same time, in violation of the Golden Bull, elections took place in the presence of the emperor [17].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The coronation took place on April 9 in Aachen.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Since his election as King of Germany in Maximilian, he has been actively involved in the administration of the empire.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

At the same time, he had to fight against the King of France, who formed an alliance against the empire with the new King Henry VII of England, as well as with King Matthias Corvin of Hungary, an enemy of the Habsburgs.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

After the death of his father on August 19, 1493, Maximilian took full power in the Empire.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

By this time, the system of governing the empire was in a deep crisis: Germany by this time had formed several hundred state entities of various levels of independence and various financial and military potential, and the emperor's levers of influence on the princes of the empire turned out to be outdated and ineffective.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Major principalities pursued a virtually independent foreign policy, while trying to subordinate the neighboring possessions of knights and imperial cities, which formed the basis of the armed forces and budget of the empire.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Frederick III, although he admired his son, often disagreements arose between them.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Thus, Friedrich categorically opposed the reforms that Maximilian tried to carry out [18].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In 1495, Maximilian I convened the General Reichstag of the Holy Roman Empire in Worms, for approval of which he submitted a project for the reform of the state administration of the empire.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

As a result of the discussion, the so-called “Imperial Reform” (German: Reichsreform) was adopted.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Germany was divided into six imperial districts (four more were added in 1512).


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The governing body of the district was the district assembly, in which all state entities in the district had the right to participate: secular and spiritual principalities, imperial knights and free cities.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Each state entity had one vote (in some districts this ensured the predominance of imperial knights, small principalities and cities that formed the main support of the emperor).


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Districts dealt with military construction, defense organization, army recruitment, and the distribution and collection of imperial taxes.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Of great importance was also the creation of the Supreme Imperial Court, the supreme judicial body of Germany, which became one of the main tools of the emperor's influence on territorial princes and a mechanism conducting a unified policy in all state entities of the empire [19] [20].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

However, Maximilian's attempts to deepen the reform of the empire and create unified executive bodies, as well as a single imperial army, failed: the princes of the empire strongly opposed and did not allow to pass through the Reichstag these proposals of the emperor.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Moreover, the imperial estates refused to finance his Italian campaigns (see below), which sharply weakened the emperor's position in the international arena and in the empire itself.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Realizing the institutional weakness of imperial power in Germany, Maximilian I continued the policy of his predecessors to separate the Austrian monarchy from the empire: relying on the Privilegium Maius 1453, as Archduke of Austria, he refused to participate in the financing of imperial institutions and did not allow imperial taxes to be levied on Austrian lands.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The duchies of Austria did not participate in the work of the imperial Reichstag and other general bodies.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Austria was effectively placed outside the empire, and its independence was expanded.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Almost all the policy of Maximilian I was carried out primarily in the interests of Austria and the Habsburg dynasty, and only secondly, Germany [20] [21].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Of great importance for the Constitution of the Holy Roman Empire was also the rejection of the principle of the need for the coronation of the emperor by the Pope in order to legitimize his rights to the title of emperor.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In 1508, he tried to make an expedition to Rome for his coronation, but was not allowed to pass by the Venetians, who controlled the routes from Germany to Italy.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

On February 4, 1508, at a festive ceremony in Trient, he was proclaimed emperor.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Pope Julius II, who urgently needed Maximilian I to create a broad coalition against Venice, allowed him to use the title of “elected emperor”.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In the future, the successors of Maximilian I (except Charles V) no longer sought coronation, and imperial law included the provision that the very election of the German king by the Electors makes him emperor [22] [23].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Italian wars


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian I, unlike his predecessors, did not leave claims to Italy.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

On August 24, 1493, Maximilian signed a marriage contract in Innsbruck with Bianca Maria Sforza, daughter of Galeazzo Maria Sforza, Duke of Milan, niece and potential heiress of Ludovico Moro, de facto ruler of Milan.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The ceremony itself took place in November 1493 in Milan (by proxy) and in March 1494 in Tyrol.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

This marriage brought him a rich dowry, but most of the money was spent on paying Maximilian's debts.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian also acquired the rights to the Duchy of Milan [24].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In March 1495, Maximilian I joined the Anti-French Holy League, which also included Spain, the Republic of Venice, the Duchy of Milan and the Papal States.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

At the same time, Maximilian managed to agree on a truce with the Republic of Venice, with which the Empire had long been in conflict over the trading cities owned by the Habsburgs on the Adriatic coast, which Venice tried to annex.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

However, as a result, in 1494, Italy was invaded by the army of King Charles VIII of France, which captured the Kingdom of Naples with almost no resistance.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

This marked the beginning of the long Italian Wars, which determined the policies of the main European states for almost half a century.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

But Charles did not manage to gain a foothold in Italy, as the League army managed to expel Charles VIII from Italy [25].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

However, already in 1499, the French, having enlisted the neutrality of Philippe the Beautiful, reentered Italy and occupied Milan.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian at that time was absorbed in the war with the Swiss Alliance, which, however, ended extremely unsuccessfully: in the Battle of Dornach, imperial troops were completely defeated by the Swiss and throughout the Basel Peace In 1499, the emperor actually recognized the independence of Switzerland not only from the Habsburgs, but also from the empire [26].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

After the Swiss Wars, Maximilian I provided military assistance to Duke Ludovico Moro, who managed to recapture Milan in 1500.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

But soon the French defeated the Milanese army, captured Ludovico and firmly established themselves in Lombardy.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

French King Louis XII was able to bring the emperor out of the struggle for Italy for some time, concluding an agreement with him on the marriage of the grandson of Maximilian Charles and the daughter of King Claude of France, promising as dowry Duchy of Burgundy and Milan.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In April 1505, Maximilian granted Louis XII an investment in the Duchy of Milan.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The Franco-Imperial Alliance was dissolved on the French initiative in 1506, but at first it did not come to an armed clash.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

In 1508, the troops of the Republic of Venice did not allow the emperor to be coronated in Rome and forced Maximilian to cede a number of territories in Istria and Friuli to Venice.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The aggressiveness of the republic led to the formation in the same year of the Antivenetian League of Cambrai, led by Pope Julius II and the King of France, which was joined by the emperor.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The French army won a number of high-profile victories over the forces of Venice in 1509, but this alarmed the Pope, who, together with Maximilian and Venice in 1510, organized the Holy League against France [27 ].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The emperor's participation in subsequent wars was extremely limited: the imperial princes refused to finance his military campaigns, and his own income from the Habsburg lands was not enough to organize large military units.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian I tried to raise funds in other ways: he laid the silver mines of Tyrol to the Fugger banking house, which later played an extremely negative role in the economic development of the region, and also actively resorted to loans from the English king.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Despite financial difficulties, it was Maximilian I who became the creator of a new type of army: the Landsknecht mercenary troops, which replaced the knightly militia and soon became the main military force in all European countries.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

His name is also associated with the beginning of an active trade in German soldiers, provided to richer foreign countries for a fee.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

At the same time, the military campaigns of the emperor himself were extremely unsuccessful: in 1511, he unsuccessfully tried to organize resistance to the French offensive in Artois, and in 1515 he undertook a campaign to Milan, which failed on the way to the city.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

As a result, French hegemony was established in Northern Italy, and the emperor lost all influence in Italian affairs.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Humanist movement


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

During the reign of Maximilian I, the humanist movement flourished in Germany.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The ideas of Erasmus of Rotterdam and the Erfurt Circle of Humanists gained European fame.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The emperor supported the arts, sciences and new philosophical ideas.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Joachim Wadian, Stiborius, Georg Tannstätter worked at his court, and the Austrian humanist Johann Kuspinian was appointed professor at the University of Vienna.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Liberation from church ideological dogmas resulted in Martin Luther's speech in Wittenberg in 1517, which marked the beginning of the Reformation in Europe [27].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Recent years


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The death of King Ferdinand II of Aragon on January 23, 1516 allowed Charles, the eldest grandson of Maximilian, to become king of the united Spanish kingdom.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Wanting to provide Charles with the imperial crown and unable to continue the fight against France, Maximilian made peace with King Francis I.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

This also made it possible to focus on protecting against Venice.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

One of the priority areas of Maximilian's activities in recent years has been Eastern politics, as well as the planned Crusade against Turkey.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

To do this, he decided to look for an ally in the person of Grand Duke of Moscow Vasily III, to whom a close friend of the emperor, Sigismund von Herberstein, was sent as an ambassador.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Pope Leo X also appealed to Christian rulers, but except Maximilian, no one showed interest in the campaign [27].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian died on January 12, 1519 in Wels.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

His body was buried under the steps of the altar of St. George's Chapel in Neustadt, and his heart was buried by will in Bruges next to his first wife, Mary of Burgundy [28].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Personal qualities


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian always paid great attention to physical exercise, hunting and had such physical strength that she was legendary.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

He was a wonderful tournament fighter [29] and a recognized expert in tournament rules, which was reflected in the work Freydal (1512-1515) compiled under his personal leadership, containing 255 engravings with images of fights, including those with the personal participation of the emperor [30].


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Marriages and children


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

1st wife: from August 19, 1477 Mary of Burgundy (1457—1482), Duchess of Burgundy from 1477, daughter of Charles the Bold, Duke of Burgundy.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Kids:


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Philip I the Beautiful (July 22, 1478 — September 25, 1506), King of Castile (c 1504), Duke of Brabant, Limburg, Luxembourg and Geldern, Palatinate of Burgundy, Count of Artois, Flanders, Gennegau (Hainaut), Holland and Zeeland, Margrave of Namur, Titular Duke of Burgundy since 1482


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Margaret (10 January 1480 — 1 December 1530), Countess of Artois, Burgundy and Charolais from 1493, stattholder of the Spanish Netherlands from 1519; 1st husband: Infante Juan of Aragon from 1497; 2nd husband: 1501 Philibert II, Duke Savoy


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Franz (September 2/10, 1481 — December 26, 1481)


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

2nd wife: March 16, 1494 (Tyrol) Bianca Maria Sforza (1472—1510), daughter of Galeazzo Maria Sforza, Duke of Milan


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian I also had several illegitimate children with mistresses.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

From an unknown person:


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Margarita (c. 1480 — June 1537); 1st husband: c.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

1497 Johann von Hille; 2nd husband: Ludwig (died 16 April 1525), Count von Helfenstein


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

From Margarita von Edelsheim:


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Barbara von Rottal (June 29, 1500 — March 31, 1550); 1st husband: July 22, 1515 Sigmund von Dietrichstein (March 19, 1484 — May 19, 1533); 2nd husband: c.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

1535 Ulrich von Kzettritz (d. 1543); 3rd husband: c.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

1544 Balthasar von Schweinitz (d. 1572)


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Georg Piret (1504 — 4 May 1557), Bishop of Brixen 1525—1539, Bishop of Brescia 1526—1538, Archbishop of Valencia 1538—1544, Bishop of Liege from 24 June 1544


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

From Anna von Helfenstein:


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian Friedrich von Amberg (June 10, 1511 — April 21, 1533), Senior Zu Feldkirch; Wife: Elisabeth von Oettingen from 1534 (June 16, 1499 — August 31, 1553)


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Leopold (1515 — 17 September 1557), Bishop of Cordoba from 1541


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Dorothea (1516—1572), heiress of Falkenburg, Darbuy and Halem, maid of honor of Queen Mary of Hungary; husband: since November 11, 1539 Johann East Frisian (d. 6 June 1572), Senor of Falkenburg, Darbuy and Halema


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Anna; m.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Louis d'Irlemont


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Anna Margarita (ca. 1517—?) , maid of honor of Queen Mary of Hungary; husband: c.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

1545 Francois de Meulun (d. 1547), Count d'Epinay from 1514, constable of Flanders


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Elizabeth (d. 1581/1584); husband: from 1531 Louis III de la Marc (died after May 6, 1544), Comte de Rochefort


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Barbara; husband: Wolfgang Plaiss


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Cornelius (September 1507 — after 1527)


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Christoph Ferdinand (died after 1522)


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Perhaps Maximilian's illegal daughter was:


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Guelma; Husband: Rudiger von Westernach


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

ascendants


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

8. Leopold III, Duke of Austria 4.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Ernst, Duke of Austria 9.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Verde Visconti2.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Frederick III, Holy Roman Emperor 10.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Zemovit IV Plotsky 5.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Kimburg Mazowiecka11.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Alexandra Olgerdovna1.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian I12.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Juan I6.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Duarte I13.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Phillip of Lancaster 3.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Eleonora Elena Portugal.14.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Ferdinand I Fair7.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Eleanor of Aragon, Queen of Portugal 15.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Eleanor d'Albuquerque


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

See also


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Landsknecht


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Maximilian armor


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Notes


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Literature


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Thomas Schauerte.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Die Ehrenpforte für Kaiser Maximilian I.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Dürer und Altdorfer im Dienst des Herrschers (= Kunstwissenschaftliche Studien; 95), München/Berlin 2001.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Larry Silver.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

Marketing Maximilian.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

The Visual Ideology of a Holy Roman Emperor, Princeton/Oxford: Princeton University Press, 2008.


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

References


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

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Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor


Question: How many wives did Maximilian I have?

Maximilian I, Holy Roman Emperor

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Maximilian I, Holy Roman Emperor

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Category:Dukes of Geldern


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Category:Counts of Charolais


Question: Who first studied cell structure in biology?

Cage

A cell is a structural and functional elementary unit of the structure and vital activity of all organisms (except viruses and viroids - forms of life that do not have a cellular structure).


Question: Who first studied cell structure in biology?

Cage

It has its own metabolism, is capable of self-reproduction.


Question: Who first studied cell structure in biology?

Cage

An organism consisting of one cell is called a unicellular organism (many protozoa and bacteria).


Question: Who first studied cell structure in biology?

Cage

The branch of biology that studies the structure and vital functions of cells is called cytology.


Question: Who first studied cell structure in biology?

Cage

It is also common to talk about cell biology, or cell biology.


Question: Who first studied cell structure in biology?

Cage

Discovery history


Question: Who first studied cell structure in biology?

Cage

The first person to see the cells was the English scientist Robert Hooke (known for the discovery of Hooke's Law).


Question: Who first studied cell structure in biology?

Cage

In 1665, while trying to understand why the cork tree floats well, Hooke began to look at thin sections of the cork with the help of a microscope he had improved.


Question: Who first studied cell structure in biology?

Cage

He discovered that the cork was divided into many tiny cells that reminded him of honeycombs in honey bee hives, and he called these cells cells cells (cell means “cell, cell” in English).


Question: Who first studied cell structure in biology?

Cage

In 1675, the Italian physician Marcello Malpighi confirmed the cellular structure of plants, and in 1681, the English botanist Nehemia Grew.


Question: Who first studied cell structure in biology?

Cage

They began to talk about the cell as a “bubble filled with nutritious juice.”


Question: Who first studied cell structure in biology?

Cage

In 1674, Dutch master Anthony van Leeuwenhoek, using a microscope, first saw “animals” in a drop of water — moving living organisms (ciliates, amoebas, bacteria).


Question: Who first studied cell structure in biology?

Cage

Levenhoek also observed animal cells for the first time — red blood cells and spermatozoa.


Question: Who first studied cell structure in biology?

Cage

Thus, by the beginning of the 18th century, scientists knew that plants had a cellular structure under high magnification, and saw some organisms that later became known as unicellular organisms.


Question: Who first studied cell structure in biology?

Cage

In 1802-1808, the French researcher Charles-Francois Mirbel found that plants consist of tissues formed by cells.


Question: Who first studied cell structure in biology?

Cage

WELL. B.


Question: Who first studied cell structure in biology?

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Lamarck in 1809 extended Mirbel's idea of cellular structure to animal organisms.


Question: Who first studied cell structure in biology?

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In 1825, the Czech scientist J.


Question: Who first studied cell structure in biology?

Cage

Purkin discovered the nucleus of an egg of birds, and in 1839 coined the term “protoplasm”.


Question: Who first studied cell structure in biology?

Cage

In 1831, the English botanist R. Brown first described the nucleus of a plant cell, and in 1833 found that the nucleus is an obligatory organoid of a plant cell.


Question: Who first studied cell structure in biology?

Cage

Since then, the main thing in the organization of cells is not the membrane, but the content.


Question: Who first studied cell structure in biology?

Cage

Cell theory


Question: Who first studied cell structure in biology?

Cage

The cellular theory of the structure of organisms was formed in 1839 by German scientists, zoologist T. Shvanny and botanist M. Schleiden, and included three provisions.


Question: Who first studied cell structure in biology?

Cage

In 1858, Rudolf Virchow supplemented it with another provision, but there were a number of mistakes in his ideas: for example, he assumed that cells were weakly connected to each other and each existed “on its own.”


Question: Who first studied cell structure in biology?

Cage

Only later it was possible to prove the integrity of the cellular system.


Question: Who first studied cell structure in biology?

Cage

In 1878, Russian scientist I. D. Chistyakov discovered mitosis in plant cells; in 1878, V. Flemming and P. AND. Peremezhko detects mitosis in animals.


Question: Who first studied cell structure in biology?

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In 1882, V. Flemming observed meiosis in animal cells, and in 1888, E.


Question: Who first studied cell structure in biology?

Cage

Strasburger - vegetable.


Question: Who first studied cell structure in biology?

Cage

Cell theory is one of the fundamental ideas of modern biology, it has become irrefutable proof of the unity of all living things and the foundation for the development of disciplines such as embryology, histology and physiology.


Question: Who first studied cell structure in biology?

Cage

To date, the theory contains the following statements:


Question: Who first studied cell structure in biology?

Cage

A cell is an elementary unit of the structure, functioning, reproduction and development of all living organisms, there is no life outside the cell.


Question: Who first studied cell structure in biology?

Cage

A cell is an integral system containing a large number of elements connected to each other — organelles.


Question: Who first studied cell structure in biology?

Cage

Cells of different organisms are similar (homologous) in structure and basic properties and have a common origin.


Question: Who first studied cell structure in biology?

Cage

The increase in the number of cells occurs by dividing them, after their DNA replicates: the cell is from the cell.


Question: Who first studied cell structure in biology?

Cage

A multicellular organism is a system of a large number of cells combined into systems of tissues and organs interconnected by humoral and nervous regulations.


Question: Who first studied cell structure in biology?

Cage

Cells of multicellular organisms have the same full fund of the genetic material of this organism, all possible potencies for the manifestation of this material, but differ in the level of expression (work) of individual genes, which leads to their morphological and functional diversity — differentiation [1].


Question: Who first studied cell structure in biology?

Cage

The number and formulation of individual provisions of modern cell theory may differ from source to source.


Question: Who first studied cell structure in biology?

Cage

Cell research methods


Question: Who first studied cell structure in biology?

Cage

For the first time, cells were seen only after the creation of optical (light) microscopes.


Question: Who first studied cell structure in biology?

Cage

Since then, microscopy has remained one of the most important methods for studying cells.


Question: Who first studied cell structure in biology?

Cage

Light microscopy, despite its low resolution, made it possible to observe living cells.


Question: Who first studied cell structure in biology?

Cage

In the 20th century, electron microscopy was invented, which made it possible to study the ultrastructure of cells.


Question: Who first studied cell structure in biology?

Cage

Various biochemical methods are used to study the functions of cells and their parts, both preparative, for example, fractionation by differential centrifugation, and analytical.


Question: Who first studied cell structure in biology?

Cage

Cell engineering methods are used for experimental and practical purposes.


Question: Who first studied cell structure in biology?

Cage

All of these methodological approaches can be used in combination with cell culture methods.


Question: Who first studied cell structure in biology?

Cage

Optical microscopy


Question: Who first studied cell structure in biology?

Cage

In an optical microscope, the magnification of an object is achieved by a series of lenses through which light passes.


Question: Who first studied cell structure in biology?

Cage

The maximum magnification is more than 1000 times.


Question: Who first studied cell structure in biology?

Cage

Resolution is also an important characteristic — the distance between two points that are still recognized separately.


Question: Who first studied cell structure in biology?

Cage

Resolution characterizes the sharpness of the image.


Question: Who first studied cell structure in biology?

Cage

This value is limited by the wavelength of the light, and even with the shortest wavelength light, ultraviolet light, it is possible to achieve a resolution of only about 200 nm; this resolution was obtained back in the late 19th century.


Question: Who first studied cell structure in biology?

Cage

The smallest structures that can be observed under an optical microscope are mitochondria and bacteria.


Question: Who first studied cell structure in biology?

Cage

Their linear size is approximately 500 nm.


Question: Who first studied cell structure in biology?

Cage

However, objects smaller than 200 nm are visible in a light microscope if they emit light themselves.


Question: Who first studied cell structure in biology?

Cage

This feature is used in fluorescence microscopy, when cellular structures or individual proteins bind to special fluorescent proteins or antibodies with fluorescent tags.


Question: Who first studied cell structure in biology?

Cage

The quality of an image obtained with an optical microscope is also affected by contrast — it can be increased using various methods of coloring cells.


Question: Who first studied cell structure in biology?

Cage

Phase-contrast, differential interference-contrast and dark-field microscopy are used to study living cells.


Question: Who first studied cell structure in biology?

Cage

Confocal microscopes improve the quality of fluorescent images [2] [3].


Question: Who first studied cell structure in biology?

Cage

Optical microscopy images


Question: Who first studied cell structure in biology?

Cage

Electron microscopy


Question: Who first studied cell structure in biology?

Cage

In the 30s of the XX century, an electron microscope was designed, in which a beam of electrons is passed through an object instead of light.


Question: Who first studied cell structure in biology?

Cage

The theoretical resolution limit for modern electron microscopes is about 0.002 nm, but for practical reasons, only about 2 nm is achieved for biological objects.


Question: Who first studied cell structure in biology?

Cage

An electron microscope can be used to study the ultrastructure of cells.


Question: Who first studied cell structure in biology?

Cage

There are two main types of electron microscopy: scanning and transmission.


Question: Who first studied cell structure in biology?

Cage

Scanning (raster) electron microscopy (SEM) is used to study the surface of an object.


Question: Who first studied cell structure in biology?

Cage

Samples are often covered with a thin film of gold.


Question: Who first studied cell structure in biology?

Cage

SEM allows you to obtain three-dimensional images.


Question: Who first studied cell structure in biology?

Cage

Transmission (transmission) electron microscopy (TEM) is used to study the internal structure of a cell.


Question: Who first studied cell structure in biology?

Cage

An electron beam is passed through an object pretreated with heavy metals that accumulate in certain structures, increasing their electron density.


Question: Who first studied cell structure in biology?

Cage

Electrons are scattered in areas of the cell with higher electron density, as a result of which these areas appear darker in images [2] [3].


Question: Who first studied cell structure in biology?

Cage

Cell fractionation


Question: Who first studied cell structure in biology?

Cage

To establish the functions of individual components of the cell, it is important to isolate them in their pure form, most often this is done using the method of differential centrifugation.


Question: Who first studied cell structure in biology?

Cage

Methods have been developed that make it possible to obtain pure fractions of any cellular organelles.


Question: Who first studied cell structure in biology?

Cage

The production of fractions begins with the destruction of the plasmalemma and the formation of a cell homogenate.


Question: Who first studied cell structure in biology?

Cage

The homogenate is sequentially centrifuged at different rates, at the first stage four fractions can be obtained: (1) nuclei and large cell fragments, (2) mitochondria, plastids, lysosomes and peroxisomes, (3) microsomes - bubbles of the Golgi apparatus and endoplasmic reticulum, (4) ribosomes, proteins and smaller molecules will remain in the supernatant.


Question: Who first studied cell structure in biology?

Cage

Further differential centrifugation of each of the mixed fractions makes it possible to obtain pure organelle preparations, to which a variety of biochemical and microscopic methods can be applied [1].


Question: Who first studied cell structure in biology?

Cage

Cell structure


Question: Who first studied cell structure in biology?

Cage

All cellular forms of life on Earth can be divided into two superkingdoms based on the structure of their constituent cells:


Question: Who first studied cell structure in biology?

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prokaryotes (pre-nuclear) - simpler in structure, arose in the process of evolution earlier;


Question: Who first studied cell structure in biology?

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eukaryotes (nuclear) - more complex, appeared later.


Question: Who first studied cell structure in biology?

Cage

The cells that make up the human body are eukaryotic.


Question: Who first studied cell structure in biology?

Cage

Despite the variety of forms, the organization of cells of all living organisms is subject to uniform structural principles.


Question: Who first studied cell structure in biology?

Cage

The cell contents are separated from the environment by a plasma membrane, or plasmalemma.


Question: Who first studied cell structure in biology?

Cage

Inside, the cell is filled with a cytoplasm, in which various organelles and cellular inclusions are located, as well as genetic material in the form of a DNA molecule.


Question: Who first studied cell structure in biology?

Cage

Each of the organelles of the cell performs its own special function, and together they all determine the vital activity of the cell as a whole.


Question: Who first studied cell structure in biology?

Cage

Prokaryotic cell


Question: Who first studied cell structure in biology?

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Prokaryotes (from Ancient Greek: prr — 'before', 'to' and — 'nucleus') are organisms that do not have, unlike eukaryotes, a formed cell nucleus and other internal membrane organoids (with the exception of flat cisterns in photosynthetic species, for example, cyanobacteria).


Question: Who first studied cell structure in biology?

Cage

The only large ring (linear in some species) double-stranded DNA molecule that contains the bulk of the genetic material of the cell (the so-called nucleoid) does not form a complex with histone proteins (so-called chromatin).


Question: Who first studied cell structure in biology?

Cage

Prokaryotes include bacteria, including cyanobacteria (blue-green algae), and archaea.


Question: Who first studied cell structure in biology?

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The descendants of prokaryotic cells are organelles of eukaryotic cells - mitochondria and plastids.


Question: Who first studied cell structure in biology?

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The main content of the cell, which fills its entire volume, is a viscous granular cytoplasm.


Question: Who first studied cell structure in biology?

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Eukaryotic cell


Question: Who first studied cell structure in biology?

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Eukaryotes (eukaryotes, from Ancient Greek: ε' 'good', 'completely' and — 'nucleus') are organisms that, unlike prokaryotes, have a formed cell nucleus delimited from the cytoplasm by a nuclear membrane.


Question: Who first studied cell structure in biology?

Cage

The genetic material is contained in several linear double-stranded DNA molecules (depending on the type of organisms, their number per nucleus can range from two to several hundred), attached from the inside to the membrane of the cell nucleus and forming in the vast majority (except dinoflagellates) a complex with histone proteins, called chromatin.


Question: Who first studied cell structure in biology?

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Eukaryote cells have a system of internal membranes that form, in addition to the nucleus, a number of other organoids (endoplasmic network, Golgi apparatus, etc.).


Question: Who first studied cell structure in biology?

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In addition, the vast majority have permanent intracellular symbionts-prokaryotes - mitochondria, and algae and plants also have plastids.


Question: Who first studied cell structure in biology?

Cage

Prokaryotic cell structure


Question: Who first studied cell structure in biology?

Cage

The cells of the two main groups of prokaryotes - bacteria and archaea - are similar in structure, their characteristic features are the absence of a nucleus and membrane organelles.


Question: Who first studied cell structure in biology?

Cage

The main components of a prokaryotic cell are:


Question: Who first studied cell structure in biology?

Cage

The cell wall that surrounds the cell from the outside protects it, gives it a stable shape that prevents osmotic destruction.


Question: Who first studied cell structure in biology?

Cage

In bacteria, the cell wall consists of peptidoglycan (murein), built from long polysaccharide chains interconnected by short peptide bridges.


Question: Who first studied cell structure in biology?

Cage

According to the structure of the cell wall, two groups of bacteria are distinguished:


Question: Who first studied cell structure in biology?

Cage

Gram-positive bacteria (for example, the genera Staphylococcus, Bacillus, Lactobacillus [4]) have a simpler cell wall structure, consisting almost exclusively of murein;


Question: Who first studied cell structure in biology?

Cage

Gram-negative bacteria (e.g. Salmonella, Escherichia, Azotobacter genera [4]) — the cell wall contains less peptidoglycan and has an additional outer membrane that consists of phospholipids .


Question: Who first studied cell structure in biology?

Cage

The archaea cell wall does not contain murein, but is built mainly from a variety of proteins and polysaccharides [3].


Question: Who first studied cell structure in biology?

Cage

The capsule is a mucous membrane found in some bacteria, located outside the cell wall.


Question: Who first studied cell structure in biology?

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It consists mainly of a variety of proteins, carbohydrates and uronic acids.


Question: Who first studied cell structure in biology?

Cage

Capsules protect cells from drying out, can help bacteria in colonies hold together, and individual bacteria attach to various substrates.


Question: Who first studied cell structure in biology?

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In addition, capsules provide the cell with additional protection: for example, encapsulated strains of pneumococci multiply freely in the body and cause pneumonia, while unencapsulated are quickly destroyed by the immune system and are completely harmless [4].


Question: Who first studied cell structure in biology?

Cage

Drank or villi are thin hair-like outgrowths that are present on the surface of bacterial cells.


Question: Who first studied cell structure in biology?

Cage

There are various types of saws, of which the most common are:


Question: Who first studied cell structure in biology?

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Fimbrias - drank, which serve to attach.


Question: Who first studied cell structure in biology?

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For example, the causative agent of gonorrhea, Neisseria gonorrhoeae, uses fimbria to retain it on the host's mucosa.


Question: Who first studied cell structure in biology?

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Sexual drinks (F-drinks) are involved in the process of conjugation in bacteria [3].


Question: Who first studied cell structure in biology?

Cage

Flagella is the organelles of the movement of some bacteria.


Question: Who first studied cell structure in biology?

Cage

The bacterial flagellum is much simpler than eukaryotic, and it is 10 times thinner, is not covered by a plasma membrane and consists of identical protein molecules that form a cylinder.


Question: Who first studied cell structure in biology?

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In the membrane, the flagellum is fixed with the help of the basal body [3].


Question: Who first studied cell structure in biology?

Cage

Plasma and internal membranes.


Question: Who first studied cell structure in biology?

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The general principle of the device of cell membranes does not differ from eukaryotes, however, there are many differences in the chemical composition of the membrane, in particular, there are no cholesterol molecules in prokaryote membranes and some lipids inherent in eukaryote membranes.


Question: Who first studied cell structure in biology?

Cage

Most prokaryotic cells (unlike eukaryotic cells) do not have internal membranes that divide the cytoplasm into separate compartments.


Question: Who first studied cell structure in biology?

Cage

Only in some photosynthetic and aerobic bacteria plasmalemma forms a bend into the cell, which performs the corresponding metabolic functions [3].


Question: Who first studied cell structure in biology?

Cage

A nucleoid is a part of the cytoplasm not bounded by membranes, in which the ring DNA molecule, the “bacterial chromosome”, is located, where all the genetic material of the cell is stored [3].


Question: Who first studied cell structure in biology?

Cage

Plasmids are small additional ring DNA molecules that usually carry only a few genes.


Question: Who first studied cell structure in biology?

Cage

Plasmids, unlike the bacterial chromosome, are not an obligatory component of the cell.


Question: Who first studied cell structure in biology?

Cage

They usually give bacteria certain beneficial properties, such as antibiotic resistance, the ability to absorb certain energy substrates from the environment, the ability to initiate sexual process, etc. [3] [4]


Question: Who first studied cell structure in biology?

Cage

Prokaryote ribosomes, like all other living organisms, are responsible for the translation process (one of the stages of protein biosynthesis).


Question: Who first studied cell structure in biology?

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However, bacterial ribosomes are slightly smaller than eukaryotic ribosomes (sedimentation coefficients 70S and 80S, respectively), and have a different composition of proteins and RNA.


Question: Who first studied cell structure in biology?

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Because of this, bacteria, unlike eukaryotes, are sensitive to antibiotics such as erythromycin and tetracycline, which selectively act on 70S ribosomes [4].


Question: Who first studied cell structure in biology?

Cage

Endospores are structures surrounded by a dense membrane that contain bacterial DNA and ensure survival in adverse conditions.


Question: Who first studied cell structure in biology?

Cage

Only some types of prokaryotes are capable of endospore formation, for example, representatives of the genus Clostridium (C. tetani is the causative agent of tetanus, C. botulinum is the causative agent of botulism, C. perfringens - the causative agent of gas gangrene, etc.) and Bacillus (in particular B. anthracis is the causative agent of anthrax).


Question: Who first studied cell structure in biology?

Cage

To form an endospore, the cell replicates its DNA and surrounds the copy with a dense membrane, removing excess water from the created structure, and slows down metabolism in it [3].


Question: Who first studied cell structure in biology?

Cage

Bacterial spores can withstand rather harsh environmental conditions, such as prolonged drying, boiling, short-wave irradiation, etc.


Question: Who first studied cell structure in biology?

Cage


Question: Who first studied cell structure in biology?

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The structure of the eukaryotic cell


Question: Who first studied cell structure in biology?

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Error: Must specify an image in the first line.


Question: Who first studied cell structure in biology?

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Animal cell surface complex


Question: Who first studied cell structure in biology?

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It consists of glycocalix, plasmalemma and the cortical layer of the cytoplasm located under it.


Question: Who first studied cell structure in biology?

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The plasma membrane is also called plasmalemma, outer cell membrane, cytolemma, etc. It's about 10 nanometers thick.


Question: Who first studied cell structure in biology?

Cage

It ensures the separation of the cell and the external environment, as well as the passage of certain substances inside and out.


Question: Who first studied cell structure in biology?

Cage

The cell does not spend energy to preserve the integrity of its membrane: molecules are held according to the same principle by which fat molecules are held together — the hydrophobic parts of the molecules thermodynamically it is more advantageous to be located in close proximity to each other.


Question: Who first studied cell structure in biology?

Cage

Glycocalyx is a molecule of oligosaccharides, polysaccharides, glycoproteins and glycolipids “anchored” in the plasmalemma.


Question: Who first studied cell structure in biology?

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Glycocalyx performs receptor and marker functions.


Question: Who first studied cell structure in biology?

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The plasma membrane of animal cells mainly consists of phospholipids and lipoproteins with protein molecules interspersed with it, in particular, surface antigens and receptors.


Question: Who first studied cell structure in biology?

Cage

In the cortical (adjacent to the plasma membrane) layer of the cytoplasm there are specific elements of the cytoskeleton — actin microfilaments ordered in a certain way.


Question: Who first studied cell structure in biology?

Cage

The main and most important function of the cortical layer (cortex) is pseudopodial reactions: ejection, attachment and contraction of pseudopodia.


Question: Who first studied cell structure in biology?

Cage

At the same time, microfilaments are rebuilt, lengthened or shortened.


Question: Who first studied cell structure in biology?

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The shape of the cell (for example, the presence of microvilli) also depends on the structure of the cytoskeleton of the cortical layer.


Question: Who first studied cell structure in biology?

Cage

Cytoplasm structure


Question: Who first studied cell structure in biology?

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The liquid component of the cytoplasm is also called cytosol.


Question: Who first studied cell structure in biology?

Cage

Under a light microscope, it seemed that the cell was filled with something like liquid plasma or sol, in which the nucleus and other organelles “float”.


Question: Who first studied cell structure in biology?

Cage

In fact, this is not the case.


Question: Who first studied cell structure in biology?

Cage

The internal space of the eukaryotic cell is strictly ordered.


Question: Who first studied cell structure in biology?

Cage

The movement of organoids is coordinated with the help of specialized transport systems, the so-called microtubules, which serve as intracellular “roads”, and special proteins of dyneins and kinesins, which play the role of “engines”.


Question: Who first studied cell structure in biology?

Cage

Individual protein molecules also do not diffuse freely throughout the intracellular space, but are sent to the necessary compartments using special signals on their surface, recognizable transport systems of the cage.


Question: Who first studied cell structure in biology?

Cage

Ribosomes


Question: Who first studied cell structure in biology?

Cage

Ribosomes are organoids that a cell needs for protein synthesis.


Question: Who first studied cell structure in biology?

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Their size is approximately 20—30 nm.


Question: Who first studied cell structure in biology?

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There are several million of them in the cage.


Question: Who first studied cell structure in biology?

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Ribosomes are formed from two subunits: large and small, consisting of four RNA molecules and several protein molecules.


Question: Who first studied cell structure in biology?

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In eukaryotic cells, ribosomes are found not only in the cytoplasm, but also in mitochondria and chloroplasts.


Question: Who first studied cell structure in biology?

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Ribosomes form in the nucleolus region, and then enter the cytoplasm through the nuclear pores.


Question: Who first studied cell structure in biology?

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Endoplasmic reticulum


Question: Who first studied cell structure in biology?

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In the eukaryotic cell, there is a system of intersecting membrane compartments (tubes, sacs and flat cisterns of different sizes) called the endoplasmic reticulum (or endoplasmic network, EPR or EPS).


Question: Who first studied cell structure in biology?

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That part of the EPR, to the membranes of which ribosomes are attached, is referred to as a granular (or rough) endoplasmic reticulum, and proteins are synthesized on its membranes.


Question: Who first studied cell structure in biology?

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Those compartments on the walls of which there are no ribosomes are referred to as agranular (or smooth) EPR, which takes part in the synthesis of lipids.


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The inner spaces of the smooth and granular EPS are not isolated, but pass into each other and communicate with the lumen of the nuclear shell.


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Golgi apparatus


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The Golgi apparatus is a stack of flat membrane tanks, slightly expanded closer to the edges.


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Some proteins synthesized on granular EPR membranes and intended for the secretion or formation of lysosomes mature in the tanks of the Golgi apparatus.


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The Golgi apparatus is asymmetrical - tanks located closer to the nucleus of the cell (cis Golgi) contain the least mature proteins, membrane vesicles - vesicles - are continuously attached to these tanks, budding off from the endoplasmic reticulum.


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Apparently, with the help of the same bubbles, the ripening proteins continue to move from one tank to another.


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In the end, bubbles containing fully mature proteins bud off from the opposite end of the organelle (Trans-Golgi).


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kernel


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The cell nucleus contains DNA molecules on which the body's genetic information is recorded.


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In the nucleus, replication takes place — doubling of DNA molecules, as well as transcription — synthesis of RNA molecules on the DNA matrix.


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In the nucleus, the synthesized RNA molecules undergo some modifications (for example, during the splicing process, insignificant, meaningless areas are excluded from messenger RNA molecules), and then go into cytoplasm.


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The assembly of ribosomes also takes place in the nucleus, in special formations called nucleoli.


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The compartment for the nucleus, the karyoteka, is formed due to the expansion and fusion of the cisterns of the endoplasmic network with each other in such a way that the nucleus has formed double walls due to the narrow ones surrounding it compartments of the nuclear envelope.


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The cavity of the nuclear shell is called the lumen or perinuclear space.


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The inner surface of the nuclear shell is underlaid by a nuclear lamine, a rigid protein structure formed by lamine proteins, to which chromosomal DNA strands are attached.


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In some places, the inner and outer membranes of the nuclear membrane merge and form so-called nuclear pores, through which material exchange between the nucleus and the cytoplasm takes place.


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Lysosomes


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Lysosome is a small body bounded from the cytoplasm by a single membrane and looks like a vesicle.


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It contains lytic enzymes that can break down almost any natural organic compounds.


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The main function is autolysis — that is, the cleavage of individual organoids, sections of the cell cytoplasm.


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Cytoskeleton


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The elements of the cytoskeleton include protein fibrillar structures located in the cytoplasm of the cell: microtubules, actin and intermediate filaments.


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Microtubules take part in the transport of organelles, are part of the organoids of movement - cilia and flagella, characteristic of some cells (for example, ciliates, spermatozoa), from microtubules a mitotic fission spindle is being built.


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Actin filaments are necessary to maintain the shape of the cell, pseudopodial reactions.


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The role of intermediate filaments also seems to be to maintain cell structure.


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Cytoskeleton proteins account for several tens of percent of the cellular protein mass.


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Centrioles


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Centrioles are cylindrical protein structures located near the nucleus of animal cells (plants do not have centrioles, with the exception of lower algae).


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The centriole is a cylinder, the lateral surface of which is formed by nine sets of microtubules.


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The number of microtubules in the kit can vary from 1 to 3 for different organisms.


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Around the centrioles there is the so-called center of organization of the cytoskeleton, the area in which the negative ends of the cell microtubules are grouped.


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Before dividing, the cell contains two centrioles located at right angles to each other.


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During mitosis, they diverge to different ends of the cell, forming the poles of the fission spindle.


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After cytokinesis, each daughter cell receives one centriole, which doubles to the next division.


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The doubling of centrioles does not occur by division, but by synthesizing a new structure perpendicular to the existing one.


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Centrioles appear to be homologous to the basal bodies of flagella and cilia.


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Mitochondria


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Mitochondria are special cell organelles, the main function of which is the synthesis of ATP, a universal energy carrier.


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Breathing (oxygen absorption and carbon dioxide release) also occurs due to mitochondrial enzymatic systems.


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The internal lumen of mitochondria, called the matrix, is delimited from the cytoplasm by two membranes, outer and inner, between which the intermembrane space is located.


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The inner membrane of mitochondria forms folds - crists, on which enzymes are placed that accelerate the oxidation reactions of fats and carbohydrates.


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The matrix contains various enzymes that are involved in the respiration and synthesis of ATP.


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The hydrogen potential of the inner membrane of mitochondria is of central importance for the synthesis of ATP.


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Mitochondria have their own DNA genome and prokaryotic ribosomes, which certainly indicates the symbiotic origin of these organelles.


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Not all mitochondrial proteins are encoded in mitochondrial DNA, most of the mitochondrial protein genes are in the nuclear genome, and their corresponding products are synthesized in the cytoplasm, and then transported to mitochondria.


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Mitochondrial genomes differ in size: for example, the genome of human mitochondria contains only 13 genes.


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The simplest Reclinomonas americana has the largest number of mitochondrial genes (97) from the studied organisms.


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Comparison of prokaryotic and eukaryotic cells


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The most important difference between eukaryotes and prokaryotes has long been the presence of a formed nucleus and membrane organoids.


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However, by the 1970s and 1980s, it became clear that this was only a consequence of deeper differences in the organization of the cytoskeleton.


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For some time, it was believed that the cytoskeleton is characteristic only of eukaryotes, but in the mid-1990s, proteins homologous to the main proteins of the eukaryote cytoskeleton were also found in bacteria.


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It is the presence of a specifically arranged cytoskeleton that allows eukaryotes to create a system of mobile internal membrane organoids.


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In addition, the cytoskeleton allows endo- and exocytosis (it is assumed that it was due to endocytosis that intracellular symbionts, including mitochondria and plastids).


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Another important function of the eukaryote cytoskeleton is to ensure the division of the nucleus (mitosis and meiosis) and the body (cytotomy) of the eukaryotic cell (the division of prokaryotic cells is easier).


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Differences in the structure of the cytoskeleton explain other differences between pro- and eukaryotes - for example, the constancy and simplicity of the forms of prokaryotic cells and the significant variety of shapes and the ability to change it eukaryotic, as well as the relatively large size of the latter.


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Thus, the size of prokaryotic cells is on average 0.5—5 microns, the size of eukaryotic cells is on average from 10 to 50 microns.


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In addition, only among eukaryotes there are truly giant cells, such as massive eggs of sharks or ostriches (in a bird's egg, the entire yolk is one huge egg), neurons of large mammals, whose processes, reinforced by the cytoskeleton, can reach tens of centimeters in length.


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Anaplasia


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The destruction of the cellular structure (for example, in malignant tumors) is called anaplasia.


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Intercellular contacts


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In higher animals and plants, cells are combined into tissues and organs, in which they interact with each other, in particular, due to direct physical contacts.


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In plant tissues, individual cells are connected to each other by means of plasmodesms, and animals form various types of cell contacts.


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Plant plasmodesms are thin cytoplasmic channels that pass through the cell walls of neighboring cells, connecting them together.


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The plasmodesm cavity is covered with plasmalemma.


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The totality of all cells united by plasmodesms is called symplast, and regulated transport of substances is possible between them.


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Intercellular contacts of vertebrates based on structure and functions are divided into three main types: anchor junctions, including adhesion contacts and desmosomes, dense or insulating (English: tight junction) and slotted or communication (English: gap junction).


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In addition, some special types of compounds between cells, such as chemical synapses of the nervous system and immunological synapses (between T-lymphocytes and antigen-presenting cells), are combined functional feature into a separate group: contacts that transmit signals (English: signal-relaying junction).


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However, anchor, slit and tight contacts can also participate in intercellular signaling [2].


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Cell cycle


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Cell division


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Eukaryotic cell division


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Amitosis is a direct division of a cell that occurs in somatic cells of eukaryotes less often than mitosis.


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In most cases, amitosis is observed in cells with reduced mitotic activity: these are aging or pathologically altered cells, often doomed to death (germ membrane cells mammals, tumor cells and others).


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In amitosis, the interphase state of the nucleus is morphologically preserved, the nucleolus and nuclear membrane are clearly visible.


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There is no DNA replication.


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Chromatin spiralization does not occur, chromosomes are not detected.


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The cell retains its characteristic functional activity, which almost completely disappears during mitosis.


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This is, for example, the division of macronucleuses of many ciliates, where short fragments of chromosomes are segregated without the formation of a spindle.


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In amitosis, only the nucleus divides, and without the formation of a fission spindle, so the hereditary material is distributed randomly.


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The absence of cytokinesis leads to the formation of binucleated cells that are not able to enter the normal mitotic cycle in the future.


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With repeated amitoses, multinucleated cells can form.


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Mitosis (from Greek: mitos - thread) is indirect cell division, the most common method of reproduction of eukaryotic cells, one of the fundamental processes of ontogenesis.


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Mitotic division ensures the growth of multicellular eukaryotes by increasing the population of tissue cells.


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The biological significance of mitosis lies in the strictly equal distribution of chromosomes between daughter nuclei, which ensures the formation of genetically identical daughter cells and preserves continuity among cell generations [9].


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The crushing of a fertilized egg and the growth of most tissues in animals also occur through mitotic divisions [10].


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On the basis of morphological features, mitosis is conditionally divided into:


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prophase,


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prometaphase


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metaphase,


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anaphase


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telophase.


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Mitosis lasts 1-2 hours on average [9] [11].


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In animal cells, mitosis usually lasts 30—60 minutes, and in plant cells it lasts 2—3 hours [12].


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Over 70 years, human cells undergo a total of about 1014 divisions [13].


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Meiosis (from Ancient Greek: decrease) or reduction cell division is the division of the nucleus of a eukaryotic cell with a decrease in the number of chromosomes by half.


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It takes place in two stages (reduction and equative stages of meiosis).


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Meiosis should not be confused with gametogenesis — the formation of specialized germ cells or gametes from undifferentiated stem cells.


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A decrease in the number of chromosomes as a result of meiosis in the life cycle leads to a transition from the diploid phase to the haploid phase.


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The restoration of ploidy (the transition from the haploid phase to the diploid phase) occurs as a result of the sexual process.


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Due to the fact that in the prophase of the first, reduction, stage there is a pairwise fusion (conjugation) of homologous chromosomes, the correct course of meiosis is possible only in diploid cells or in even polyploids (tetra-, hexaploid cells, etc.).


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Meiosis can also occur in odd polyploids (tri-, pentaploid, etc.), but in them, due to the inability to ensure pairwise fusion of chromosomes in prophase I, the chromosome divergence occurs with disorders that jeopardize the viability of a cell or a multicellular haploid organism that develops from it.


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The same mechanism underlies the sterility of interspecific hybrids.


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Chromosomal mutations (large-scale deletions, duplications, inversions or translocations) also impose certain restrictions on chromosome conjugation.


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Prokaryotic cell division


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Prokaryotic cells divide in two.


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First, the cell lengthens, forms a transverse septum in it.


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At the final stage, the daughter cells diverge.


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A distinctive feature of prokaryotic cell division is the direct participation of replicated DNA in the division process [14].


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Usually, prokaryotic cells divide to form two daughter cells of the same size, which is why this process is sometimes called binary division.


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Due to the fact that in the vast majority of cases prokaryotic cells have a cell wall, binary division is accompanied by the formation of a septa, a septa between daughter cells, which then exfoliates in the middle.


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The process of prokaryotic cell division has been studied in detail using the example of Escherichia coli [15].


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Differentiation of multicellular cells


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Multicellular organisms consist of cells that differ in structure and function to varying degrees, for example, an adult has about 230 different types of cells [16].


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All of them are descendants of the same cell — the zygote (in the case of sexual reproduction) — and acquire differences as a result of the differentiation process.


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Differentiation in the vast majority of cases is not accompanied by a change in the hereditary information of the cell, but is provided only by regulating gene activity, the specific nature of expression genes are inherited during maternal cell division, usually due to epigenetic mechanisms.


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However, there are exceptions: for example, during the formation of cells of a specific immune system of vertebrates, some genes are restructured, mammalian red blood cells lose all hereditary information, and germ cells are half of it.


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Differences between cells at the first stages of embryonic development appear, firstly, due to the heterogeneity of the cytoplasm of the fertilized egg, which causes them to form during the fragmentation process cells that differ in the content of certain proteins and RNA; secondly, the microenvironment of the cell plays an important role — its contacts with other cells and the environment.


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When differentiated, cells lose their potency, that is, the ability to give rise to cells of other types.


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Totipotent cells, which include, in particular, zygote, can form an integral organism.


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Pluripotent cells (for example, blastocyst cells) have the ability to differentiate into any type of cell in the body, but extra-embryonic tissues, and therefore a new individual, cannot develop from them.


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Cells that can only give rise to a limited number of other tissues are called multipotent (adult stem cells), and those that can only reproduce themselves similar — unipotent.


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Many of the completely differentiated cells (for example, neurons, red blood cells) completely lose their ability to divide and leave the cell cycle [2].


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In some cases, differentiation may be reversed, the opposite process is called dedifferentiation.


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It is typical for regeneration processes.


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With some reservations, tumor transformation of cells can be attributed to the phenomenon of dedifferentiation [17].


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Cell death


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Single-celled organisms can in a sense be considered “immortal” because, except in cases of damage or starvation, they do not die, but divide, resulting in the formation of two new the body.


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But all cells of multicellular organisms (except gametes) are doomed to death, but they die not only if the entire individual dies - this process occurs constantly.


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The death of some cells is necessary during embryonic development, cells continue to die in adult organisms, for example, billions of cells die every hour in the human bone marrow and intestines.


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Due to physiological conditions, “programmed cell death” occurs, in other words, cells “commit suicide”.


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The most common, but not the only, method of cellular self-destruction is apoptosis.


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The main signs of apoptosis are DNA fragmentation, the decay of the cell into apoptic bodies — vesicles surrounded by membranes.


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On their surface there are special molecules that encourage neighboring cells and macrophages to phagocyte them in such a way that the process is not accompanied by inflammation.


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Apoptosis is a volatile process and requires the use of ATP.


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This path of cell death is important not only for the development of the body, the normal functioning of the immune system, but also for protecting the individual from damaged cells that may be on the way malignant transformation, and from viral infections [18].


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Physical or chemical damage to cells, as well as a lack of energy and oxygen sources, can lead to another death, necrotic death.


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Necrosis, unlike apoptosis, is a passive process, it is often accompanied by rupture of the plasmalemma and leakage of the cytoplasm.


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Necrosis almost always causes inflammation of surrounding tissues.


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Recently, the mechanism of programmed necrosis as a possible antiviral and anticancer protection has been studied [18].


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With a prolonged lack of ATP in the cell, it does not immediately die by necrosis, and in many cases it takes the path of autophagy, a process that allows it to remain for some time viable.


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With autophagy (literally “self-eating”), the metabolism switches towards active catabolism, while individual organelles are surrounded by double membranes, so-called autophagosomes are formed, merging with lysosomes, where organic substances are digested.


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If the hunger strike continues and after most of the organelles have already been “eaten”, the cell dies by necrosis.


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Some authors believe that under certain conditions autophagy can be a separate type of cell death [18].


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Cell evolution


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It is not known for certain when the first cell appeared on Earth and how it arose.


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The earliest probable fossil cell remains, estimated to be 3.49 billion years old, were found in eastern Pilbara, Australia, although the biogenicity of their origin was placed under doubt.


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Stromatolites of the same period also testify to the existence of life in early archaea [19] [20].


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The appearance of the first cells should have been preceded by the accumulation of organic substances in the environment and the appearance of a certain form of prebiotic metabolism.


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Protocells contained at least two mandatory elements: hereditary information in the form of molecules capable of self-replication, and a certain kind of shell that protected the internal contents the first cells from the environment.


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The most likely candidate for the role of self-replicative molecules is RNA, since it can simultaneously act as both a carrier of hereditary information and a catalyst; in addition, RNA, unlike from DNA, is self-sufficient for protein biosynthesis [20] [21].


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It is also unknown what substances the membranes of the first cells were built from, however, it is likely that these could be simple amphiphilic compounds, such as fatty acid salts, capable of self-organize into liposomes that can go through growth and division cycles.


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Fatty acids have been synthesized in many experiments to reproduce prebiotic conditions, and they have also been found in meteorites [21] [22].


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It is believed that the first living cells were heterotrophic.


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The emergence of eukaryotic cells


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rRNA sequencing data made it possible to build a universal tree of life, in which the last universal common ancestor gave rise to two branches of evolution: eubacteria and the Neomura clade, the last of which in turn, it was divided into two branches: archaea and eukaryotes [23].


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Endosymbiosis probably played an important role in the evolution of eukaryotes — it is believed that this is the method that nuclear cells obtained mitochondria, and later chloroplasts [24].


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Eukaryotes have many genes in common with both eubacteria and archaea; some scientists believe that they arose as a result of the fusion of the genomes of these two groups of organisms, which could have occurred as a result of endosymbiosis.


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Because of this, instead of the “tree of life”, it is proposed to use the “circle of life” [25].


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Other researchers, noting the importance of intensive horizontal transfer between the ancestors of eukaryotes, bacteria and archaeobacteria, suggest displaying phylogenetic relationships between them using a “grid life” [26].


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Chemical organization of the cell


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Group 1 (up to 98%) (organogens)


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Carbon


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Hydrogen


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Oxygen


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Nitrogen


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Group 2 (1.5—2%) (macronutrients)


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Potassium


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Sodium


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Calcium


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Magnesium


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Chlorine


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Iron


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Group 3 (>0.01%) (micronutrients)


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Zinc


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Manganese


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copper


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Fluorine


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Iodine


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Cobalt


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Molybdenum


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Group 4 (>0.00001%) (ultramicronutrients)


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Uranus


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Radium


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Gold


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See also


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Cell structure comparison in bacteria, plants, animals and fungi


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Cell chemical composition


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Cell theory


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Cell mechanics


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Cell transplantology


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Cell-free system


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Notes


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Literature


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CS1 maint: multiple names: authors list (link)


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Bilic G. L., Kryzhanovsky V. A. Biology.