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Participants reduced their greenhouse gas and water footprints by enjoying a delicious Meatless Monday on a Sunday potluck at the Resource Center for Nonviolence (rcnv.org), and then about a fourth of us began a People's Climate March through town, while passing cars honked in support!
Oogmasker met een drainerende werking.Stimuleert de microcirculatie en heeft een ontzwellende werking.
Vermindert vetmassa onder de ogen. Als masker 2 tot 3 x in de week gebruiken.
Inwerk tijd 20 minuten bij een dikkere laag. Of elke nacht in een dunnere laag als nachtverzorging.
"SUMMER 2017 DAYLIFE" at The Pool Harrah's Atlantic City Saturday July 22, 2017
The Pool After Dark inside Harrah's Resort Atlantic City, NJ | Celebrating a birthday, bachelor, or bachelorette party? For FREE or Reduced Admission, VIP, or Bottle Service to Atlantic City's top nightlife destination, get on the nightclub guest list at www.gocoastalac.com.
The Student Art Alliance of MTSU created this installation using abandoned printer paper gathered from the library recycling bins. The goal of the installation was to raise awareness and reduce paper waste. Photo by Kristen West.
I was a bit stumped as to what to do for "Reduce and Simplify" as there haven't been a huge number of elements in any of the #creativityex photographs I've done so far. Then I thought of something cheeky - I could remove the grass and leaf from my grass and leaf photo! So here is the police chalk outline of a dead leaf. Only, it turns out I only had yellow pavement chalk and I can't draw very well. I get points for creativity though, right?!
March 23, 2012. Somerville, MA.
The Centers for Disease Control and Prevention (CDC) has awarded the Massachusetts Department of Health a $1.57 million Community Transformation Grant to promote health and wellness in Middlesex County. United States Surgeon General Regina M. Benjamin, MD, MBA, formally launched the grant’s activities at a press conference at the Holiday Inn in Somerville.
The Metropolitan Area Planning Council (MAPC) collaborated with the Massachusetts Department of Public Health (MDPH) in applying for the Community Transformation Grant. This grant allows for the expansion of MDPH’s “Mass in Motion” initiative, which works to reduced obesity and supports MDPH’s smoking cessation and prevention activities.
© 2012 Marilyn Humphries
The monument preserves the ruins of the three largest of nine pueblos in this area -- Quarai, Abó, and Gran Quivira (Las Humanas). I visited Quarai and Abó.
Church building began in the 1620s but all the Salinas pueblos were abandoned during the 1670s, their populations having been reduced by forced labor, European diseases, drought, famine, and Apache attacks.
Pic by Sara Monk, shoestringonline.co.uk
This picture is one of the best of what Mrs Monk shot in London 2006
Arsehole Signs for Sale
5 December 2011 I was surprised to learn that several people had googled the word “Arsehole”, and had accordingly found this website, perhaps because of this 2006 page.
shoestringonline.co.uk/DiaryLatest/html/arsehole.html
Even in 2006 there were annoying Jeremy Clarksons everywhere.
The history of Rome includes the history of the city of Rome as well as the civilisation of ancient Rome. Roman history has been influential on the modern world, especially in the history of the Catholic Church, and Roman law has influenced many modern legal systems. Roman history can be divided into the following periods:
Pre-historical and early Rome, covering Rome's earliest inhabitants and the legend of its founding by Romulus
The period of Etruscan dominance and the regal period, in which, according to tradition, Romulus was the first of seven kings
The Roman Republic, which commenced in 509 BC when kings were replaced with rule by elected magistrates. The period was marked by vast expansion of Roman territory. During the 5th century BC, Rome gained regional dominance in Latium. With the Punic Wars from 264 to 146 BC, ancient Rome gained dominance over the Western Mediterranean, displacing Carthage as the dominant regional power.
The Roman Empire followed the Republic, which waned with the rise of Julius Caesar, and by all measures concluded after a period of civil war and the victory of Caesar's adopted son, Octavian, in 27 BC over Mark Antony.
The Western Roman Empire collapsed in 476 after the city was conquered by the Ostrogothic Kingdom. Consequently Rome's power declined, and it eventually became part of the Eastern Roman Empire, as the Duchy of Rome, from the 6th to 8th centuries. At this time, the city was reduced to a fraction of its former size, being sacked several times in the 5th to 6th centuries, even temporarily depopulated entirely.
Medieval Rome is characterised by a break with Constantinople and the formation of the Papal States. The Papacy struggled to retain influence in the emerging Holy Roman Empire, and during the saeculum obscurum, the population of Rome fell to as low as 30,000 inhabitants. Following the East–West Schism and the limited success in the Investiture Controversy, the Papacy did gain considerable influence in the High Middle Ages, but with the Avignon Papacy and the Western Schism, the city of Rome was reduced to irrelevance, its population falling below 20,000. Rome's decline into complete irrelevance during the medieval period, with the associated lack of construction activity, assured the survival of very significant ancient Roman material remains in the centre of the city, some abandoned and others continuing in use.
The Roman Renaissance occurred in the 15th century, when Rome replaced Florence as the centre of artistic and cultural influence. The Roman Renaissance was cut short abruptly with the devastation of the city in 1527, but the Papacy reasserted itself in the Counter-Reformation, and the city continued to flourish during the early modern period. Rome was annexed by Napoleon and was part of the First French Empire from 1798 to 1814.
Modern history, the period from the 19th century to the present. Rome came under siege again after the Allied invasion of Italy and was bombed several times. It was declared an open city on 14 August 1943. Rome became the capital of the Italian Republic (established in 1946). With a population of 4.4 million (as of 2015; 2.9 million within city limits), it is the largest city in Italy. It is among the largest urban areas of the European Union and classified as a global city.
Attempts have been made to find a linguistic root for the name Rome. Possibilities include derivation from the Greek Rhṓmē (Ῥώμη), meaning "bravery" or "courage"; Compare also Rumon, former name of the Tiber River. Its further etymology remains unknown, as with most Etruscan words. Thomas G. Tucker's Concise Etymological Dictionary of Latin (1931) suggests that the name is most probably from *urobsma (cf. urbs, robur) and otherwise, "but less likely" from *urosma "hill" (cf. Skt. varsman- "height, point," Old Slavonic врьхъ "top, summit", Russ. верх "top; upward direction", Lith. virsus "upper").
Rome timeline
753 BCAccording to legend, Romulus founds Rome.
753–509 BCRule of the seven Kings of Rome.
509 BCCreation of the Republic.
390 BCThe Gauls invade Rome. Rome sacked.
264–146 BCPunic Wars.
146–44 BCSocial and Civil Wars. Emergence of Marius, Sulla, Pompey and Caesar.
44 BCJulius Caesar assassinated.
There is archaeological evidence of human occupation of the Rome area from at least 5,000 years, but the dense layer of much younger debris obscures Palaeolithic and Neolithic sites. The evidence suggesting the city's ancient foundation is also obscured by the legend of Rome's beginning involving Romulus and Remus.
The traditional date for the founding of Rome is 21 April 753 BC, following M. Terentius Varro, and the city and surrounding region of Latium has continued to be inhabited with little interruption since around that time. Excavations made in 2014 have revealed a wall built long before the city's official founding year. Archaeologists uncovered a stone wall and pieces of pottery dating to the 9th century BC and the beginning of the 8th century BC, and there is evidence of people arriving on the Palatine hill as early as the 10th century BC.
The site of Sant'Omobono Area is crucial for understanding the related processes of monumentalisation, urbanisation, and state formation in Rome in the late Archaic period. The Sant'Omobono temple site dates to 7th–6th century BC, making these the oldest known temple remains in Rome.
The city's name was long credited to the legendary culture hero Romulus. It was said that Romulus and his twin brother Remus were the offspring of the rape of an Alban princess by the war god Mars and, via their mother, were further descended from the Trojan prince Aeneas, supposed son of the Greek love goddess Aphrodite. Exposed on the Tiber, they were suckled by a she-wolf and raised by a shepherd and his wife. Avenging themselves on their usurping grand-uncle and restoring their grandfather Numitor to Alba Longa's throne, they were ordered or decided to settle the hills around Rome's later Forum Boarium, an important river port connected in Roman myth with Hercules's tenth labour, capturing the cattle of Geryon.
Disputing some point of the founding or its related auguries, Remus was murdered by Romulus or one of his supporters. Romulus then established a walled and roughly square settlement, whose sacred boundary and gates were established by a ploughing ritual. Romulus then declared the town an asylum, permitted men of all classes to come to Rome as citizens, including criminals, runaway slaves, and freemen without distinction. To provide his citizens with wives, Romulus invited the neighbouring tribes to a festival in Rome where the Romans abducted many of their young women. After the ensuing war with the Sabines, Romulus shared Rome's kingship with the Sabine king Titus Tatius. Romulus selected 100 of the most noble men to form the Roman Senate, initially serving as his advisory council. These men he called fathers (Latin: patres), and their descendants became the patricians. He created three centuries of equites: Ramnes (meaning Romans), Tities (after the Sabine king), and Luceres (Etruscans). He also divided the general populace into thirty curiae, named after thirty of the Sabine women who had intervened to end the war between Romulus and Tatius. The curiae formed the voting units in the Comitia Curiata.
Rome grew from pastoral settlements on the Palatine Hill and surrounding hills approximately 30 km (19 mi) from the Tyrrhenian Sea on the south side of the Tiber. The Quirinal Hill was probably an outpost for the Sabines, another Italic-speaking people. At this location, the Tiber forms a Z-shaped curve that contains an island where the river can be forded. Because of the river and the ford, Rome was at a crossroads of traffic following the river valley and of traders travelling north and south on the west side of the peninsula.
Archaeological finds have confirmed that there were two fortified settlements in the 8th century BC, in the area of the future Rome: Rumi on the Palatine Hill, and Titientes on the Quirinal Hill, backed by the Luceres living in the nearby woods. These were simply three of numerous Italic-speaking communities that existed in Latium, a plain on the Italian peninsula, by the 1st millennium BC. The origins of the Italic peoples lie in prehistory and are therefore not precisely known, but their Indo-European languages migrated from the east in the second half of the 2nd millennium BC.
According to Dionysius of Halicarnassus, many Roman historians—including Cato and Sempronius—considered the Italian aborigines to have been prehistoric Greek colonists. The Romans then considered themselves a mix of these people, the Albans, and the other Latins, considered a blend of Pelasgians, Arcadians, Epeans, and refugee Trojans. Over time, the Etruscans and other ancient Italic peoples were admitted as citizens as well. The Sabines—considered to be Gaulish along with the other Umbri peoples of central Italy— were first mentioned in Dionysius's account for having captured the city of Lista by surprise, which was regarded as the mother-city of the Aborigines.
The Italic speakers in the area included Latins (in the west), Sabines (in the upper valley of the Tiber), Umbrians (in the north-east), Samnites (in the South), Oscans, and others. In the 8th century BC, they shared the peninsula with two other major ethnic groups: the Etruscans in the North and the Greeks in the south.
The Etruscans (Etrusci or Tusci in Latin) are attested north of Rome in Etruria (modern northern Lazio, Tuscany and part of Umbria). They founded cities such as Tarquinia, Veii, and Volterra and deeply influenced Roman culture, as clearly shown by the Etruscan origin of some of the mythical Roman kings. Historians have no literature, nor texts of religion or philosophy; therefore, much of what is known about this civilisation is derived from grave goods and tomb findings.
The Greeks had founded many colonies in Southern Italy between 750 and 550 BC (which the Romans later called Magna Graecia), such as Cumae, Naples, Reggio Calabria, Crotone, Sybaris, and Taranto, as well as in the eastern two-thirds of Sicily.
After 650 BC, the Etruscans became dominant in Italy and expanded into north-central Italy. Roman tradition claimed that Rome had been under the control of seven kings from 753 to 509 BC beginning with the mythical Romulus who was said to have founded the city of Rome along with his brother Remus. The last three kings were said to be Etruscan (at least partially)—namely Tarquinius Priscus, Servius Tullius and Tarquinius Superbus. (Priscus is said by the ancient literary sources to be the son of a Greek refugee and an Etruscan mother.) Their names refer to the Etruscan town of Tarquinia.
Livy, Plutarch, Dionysius of Halicarnassus, and others claim that Rome was ruled during its first centuries by a succession of seven kings. The traditional chronology, as codified by Varro, allots 243 years for their reigns, an average of almost 35 years, which has been generally discounted by modern scholarship since the work of Barthold Georg Niebuhr. The Gauls destroyed much of Rome's historical records when they sacked the city after the Battle of the Allia in 390 BC (according to Polybius, the battle occurred in 387/386) and what was left was eventually lost to time or theft. With no contemporary records of the kingdom existing, all accounts of the kings must be carefully questioned. The list of kings is also of dubious historical value, though the last-named kings may be historical figures. It is believed by some historians (again, this is disputed) that Rome was under the influence of the Etruscans for about a century. During this period, a bridge was built called the Pons Sublicius to replace the Tiber ford, and the Cloaca Maxima was also built; the Etruscans are said to have been great engineers of this type of structure. From a cultural and technical point of view, Etruscans had arguably the second-greatest impact on Roman development, only surpassed by the Greeks.
Expanding further south, the Etruscans came into direct contact with the Greeks and initially had success in conflicts with the Greek colonists; after which, Etruria went into a decline. Taking advantage of this, Rome rebelled and gained independence from the Etruscans around 500 BC. It also abandoned monarchy in favour of a republican system based on a Senate, composed of the nobles of the city, along with popular assemblies which ensured political participation for most of the freeborn men and elected magistrates annually.
The Etruscans left a lasting influence on Rome. The Romans learned to build temples from them, and the Etruscans may have introduced the worship of a triad of gods—Juno, Minerva, and Jupiter—from the Etruscan gods: Uni, Menrva, and Tinia. However, the influence of Etruscan people in the development of Rome is often overstated. Rome was primarily a Latin city. It never became fully Etruscan. Also, evidence shows that Romans were heavily influenced by the Greek cities in the South, mainly through trade.
The commonly held stories of the early part of the Republic (before roughly 300 BC, when Old Latin inscriptions and Greek histories about Rome provide more concrete evidence of events) are generally considered to be legendary, their historicity being a topic of debate among classicists. The Roman Republic traditionally dates from 509 BC to 27 BC. After 500 BC, Rome is said to have joined with the Latin cities in defence against incursions by the Sabines. Winning the Battle of Lake Regillus in 493 BC, Rome established again the supremacy over the Latin countries it had lost after the fall of the monarchy. After a lengthy series of struggles, this supremacy became fixed in 393, when the Romans finally subdued the Volsci and Aequi. In 394 BC, they also conquered the menacing Etruscan neighbour of Veii. The Etruscan power was now limited to Etruria itself, and Rome was the dominant city in Latium.
A formal treaty was agreed with the city-state of Carthage in 509 BC which defined the spheres of influence of each city and regulated trade between them.
Rome's early enemies were the neighbouring hill tribes of the Volscians, the Aequi, and of course the Etruscans. As years passed and military successes increased Roman territory, new adversaries appeared. The fiercest were the Gauls, a loose collective of peoples who controlled much of Northern Europe including what is modern North and Central-East Italy.
In 387 BC, Rome was sacked and burned by the Senones coming from eastern Italy and led by Brennus, who had successfully defeated the Roman army at the Battle of the Allia in Etruria. Multiple contemporary records suggest that the Senones hoped to punish Rome for violating its diplomatic neutrality in Etruria. The Senones marched 130 kilometres (81 mi) to Rome without harming the surrounding countryside; once they had sacked the city, the Senones withdrew from Rome. Brennus was defeated by the dictator Furius Camillus at Tusculum soon afterwards.
After that, Rome hastily rebuilt its buildings and went on the offensive, conquering the Etruscans and seizing territory from the Gauls in the north. After 345 BC, Rome pushed south against other Latins. Their main enemy in this quadrant were the fierce Samnites, who outsmarted and trapped the legions in 321 BC at the Battle of Caudine Forks. In spite of these and other temporary setbacks, the Romans advanced steadily. By 290 BC, Rome controlled over half of the Italian peninsula. In the 3rd century BC, Rome brought the Greek poleis in the south under its control as well.
Amidst the never-ending wars (from the beginning of the Republic up to the Principate, the doors of the temple of Janus were closed only twice—when they were open it meant that Rome was at war), Rome had to face a severe major social crisis, the Conflict of the Orders, a political struggle between the Plebeians (commoners) and Patricians (aristocrats) of the ancient Roman Republic, in which the Plebeians sought political equality with the Patricians. It played a major role in the development of the Constitution of the Roman Republic. It began in 494 BC, when, while Rome was at war with two neighbouring tribes, the Plebeians all left the city (the first Plebeian Secession). The result of this first secession was the creation of the office of Plebeian Tribune, and with it the first acquisition of real power by the Plebeians.
According to tradition, Rome became a republic in 509 BC. However, it took a few centuries for Rome to become the great city of popular imagination. By the 3rd century BC, Rome had become the pre-eminent city of the Italian peninsula. During the Punic Wars between Rome and the great Mediterranean empire of Carthage (264–146 BC), Rome's stature increased further as it became the capital of an overseas empire for the first time. Beginning in the 2nd century BC, Rome went through a significant population expansion as Italian farmers, driven from their ancestral farmlands by the advent of massive, slave-operated farms called latifundia, flocked to the city in great numbers. The victory over Carthage in the First Punic War brought the first two provinces outside the Italian peninsula, Sicily and Sardinia. Parts of Spain (Hispania) followed, and in the beginning of the 2nd century the Romans got involved in the affairs of the Greek world. By then all Hellenistic kingdoms and the Greek city-states were in decline, exhausted from endless civil wars and relying on mercenary troops.
The Romans looked upon the Greek civilisation with great admiration. The Greeks saw Rome as a useful ally in their civil strifes, and it was not long before the Roman legions were invited to intervene in Greece. In less than 50 years the whole of mainland Greece was subdued. The Roman legions crushed the Macedonian phalanx twice, in 197 and 168 BC; in 146 BC the Roman consul Lucius Mummius razed Corinth, marking the end of free Greece. The same year Cornelius Scipio Aemilianus, the son of Scipio Africanus, destroyed the city of Carthage, making it a Roman province.
In the following years, Rome continued its conquests in Spain with Tiberius Gracchus, and it set foot in Asia, when the last king of Pergamum gave his kingdom to the Roman people. The end of the 2nd century brought another threat, when a great host of Germanic peoples, namely Cimbri and Teutones, crossed the river Rhone and moved to Italy. Gaius Marius was consul five consecutive times (seven total), and won two decisive battles in 102 and 101 BC. He also reformed the Roman army, giving it such a good reorganisation that it remained unchanged for centuries.
The first thirty years of the last century BC were characterised by serious internal problems that threatened the existence of the Republic. The Social War, between Rome and its allies, and the Servile Wars (slave uprisings) were hard conflicts, all within Italy, and forced the Romans to change their policy with regards to their allies and subjects. By then Rome had become an extensive power, with great wealth which derived from the conquered people (as tribute, food or manpower, i.e. slaves). The allies of Rome felt bitter since they had fought by the side of the Romans, and yet they were not citizens and shared little in the rewards. Although they lost the war, they finally got what they asked, and by the beginning of the 1st century AD practically all free inhabitants of Italy were Roman citizens.
However, the growth of the Imperium Romanum (Roman power) created new problems, and new demands, that the old political system of the Republic, with its annually elected magistrates and its sharing of power, could not solve. Sulla's civil war and his later dictatorship, the extraordinary commands of Pompey Magnus, and the first triumvirate made that clear. In January 49 BC, Julius Caesar the conqueror of Gaul, crossed the Rubicon with his legions, occupying Rome and beginning a civil war with Pompey. In the following years, he vanquished his opponents, and ruled Rome for four years. After his assassination in 44 BC, the Senate tried to reestablish the Republic, but its champions, Marcus Junius Brutus (descendant of the founder of the republic) and Gaius Cassius Longinus were defeated by Caesar's lieutenant Marcus Antonius and Caesar's nephew, Octavian.
The years 44–31 BC mark the struggle for power between Marcus Antonius and Octavian (later known as Augustus). Finally, on 2 September 31 BC, in the Greek promontory of Actium, the final battle took place in the sea. Octavian was victorious, and became the sole ruler of Rome (and its empire). That date marks the end of the Republic and the beginning of the Principate.
Rome timeline
44 BC – AD 14Augustus establishes the Empire
AD 64Great Fire of Rome during Nero's rule
69–96Flavian dynasty; building of the Colosseum
3rd centuryCrisis of the Third Century; building of the Baths of Caracalla and the Aurelian Walls
284–337Diocletian and Constantine; building of the first Christian basilicas; Battle of Milvian Bridge; Rome is replaced by Constantinople as the capital of the Empire
395Definitive separation of Western and Eastern Roman Empire
410The Goths of Alaric sack Rome
455The Vandals of Gaiseric sack Rome
476Fall of the west empire and deposition of the final emperor Romulus Augustus
6th centuryGothic War (535–554): The Goths cut off the aqueducts in the siege of 537, an act which historians traditionally regard as the beginning of the Middle Ages in Italy
608Emperor Phocas donates the Pantheon to Pope Boniface IV, converting it into a Christian church; Column of Phocas (the last addition made to the Forum Romanum) is erected
630The Curia Julia (vacant since the disappearance of the Roman Senate) is transformed into the basilica of Sant'Adriano al Foro
663Constans II visits Rome for twelve days—the only emperor to set foot in Rome for two centuries. He strips buildings of their ornaments and bronze to be carried back to Constantinople
751Lombard conquest of the Exarchate of Ravenna; the Duchy of Rome is now completely cut off from the empire
754Alliance with the Franks; Pepin the Younger, King of the Franks, declared Patrician of the Romans, invades Italy; establishment of the Papal States
Life in Rome; animation in Latin with English subtitles
By the end of the Republic, the city of Rome had achieved a grandeur befitting the capital of an empire dominating the whole of the Mediterranean. It was, at the time, the largest city in the world. Estimates of its peak population range from 450,000 to over 3.5 million people with estimates of 1 to 2 million being most popular with historians. This grandeur increased under Augustus, who completed Caesar's projects and added many of his own, such as the Forum of Augustus and the Ara Pacis. He is said to have remarked that he found Rome a city of brick and left it a city of marble (Urbem latericium invenit, marmoream reliquit). Augustus's successors sought to emulate his success in part by adding their own contributions to the city. In AD 64, during the reign of Nero, the Great Fire of Rome left much of the city destroyed, but in many ways it was used as an excuse for new development.
Rome was a subsidised city at the time, with roughly 15 to 25 percent of its grain supply being paid by the central government. Commerce and industry played a smaller role compared to that of other cities like Alexandria. This meant that Rome had to depend upon goods and production from other parts of the Empire to sustain such a large population. This was mostly paid by taxes that were levied by the Roman government. If it had not been subsidised, Rome would have been significantly smaller.
Rome's population declined after its apex in the 2nd century. At the end of that century, during the reign of Marcus Aurelius, the Antonine Plague killed 2,000 people a day. Marcus Aurelius died in 180, his reign being the last of the "Five Good Emperors" and Pax Romana. His son Commodus, who had been co-emperor since AD 177, assumed full imperial power, which is generally associated with the beginning of the decline of the Western Roman Empire. Rome's population was only a fraction of its peak when the Aurelian Wall was completed in AD 273 (in that year its population was only around 500,000).
Starting in the early 3rd century, matters changed. The "Crisis of the Third Century" defines the disasters and political troubles for the Empire, which nearly collapsed. The new feeling of danger and the menace of barbarian invasions was clearly shown by the decision of Emperor Aurelian, who at year 273 finished encircling the capital itself with a massive wall which had a perimeter that measured close to 20 km (12 mi). Rome formally remained capital of the empire, but emperors spent less and less time there. At the end of 3rd century Diocletian's political reforms, Rome was deprived of its traditional role of administrative capital of the Empire. Later, western emperors ruled from Milan or Ravenna, or cities in Gaul. In 330, Constantine I established a second capital at Constantinople.
Christianity reached Rome during the 1st century AD. For the first two centuries of the Christian era, Imperial authorities largely viewed Christianity simply as a Jewish sect rather than a distinct religion. No emperor issued general laws against the faith or its Church, and persecutions, such as they were, were carried out under the authority of local government officials. A surviving letter from Pliny the Younger, governor of Bythinia, to the emperor Trajan describes his persecution and executions of Christians; Trajan notably responded that Pliny should not seek out Christians nor heed anonymous denunciations, but only punish open Christians who refused to recant.
Suetonius mentions in passing that during the reign of Nero "punishment was inflicted on the Christians, a class of men given to a new and mischievous superstition" (superstitionis novae ac maleficae). He gives no reason for the punishment. Tacitus reports that after the Great Fire of Rome in AD 64, some among the population held Nero responsible and that the emperor attempted to deflect blame onto the Christians. The war against the Jews during Nero's reign, which so destabilised the empire that it led to civil war and Nero's suicide, provided an additional rationale for suppression of this 'Jewish' sect.
Diocletian undertook what was to be the most severe and last major persecution of Christians, lasting from 303 to 311. Christianity had become too widespread to suppress, and in 313, the Edict of Milan made tolerance the official policy. Constantine I (sole ruler 324–337) became the first Christian emperor, and in 380 Theodosius I established Christianity as the official religion.
Under Theodosius, visits to the pagan temples were forbidden, the eternal fire in the Temple of Vesta in the Roman Forum extinguished, the Vestal Virgins disbanded, auspices and witchcraft punished. Theodosius refused to restore the Altar of Victory in the Senate House, as asked by remaining pagan Senators.
The Empire's conversion to Christianity made the Bishop of Rome (later called the Pope) the senior religious figure in the Western Empire, as officially stated in 380 by the Edict of Thessalonica. In spite of its increasingly marginal role in the Empire, Rome retained its historic prestige, and this period saw the last wave of construction activity: Constantine's predecessor Maxentius built buildings such as its basilica in the Forum, Constantine himself erected the Arch of Constantine to celebrate his victory over Maxentius, and Diocletian built the greatest baths of all. Constantine was also the first patron of official Christian buildings in the city. He donated the Lateran Palace to the Pope, and built the first great basilica, the old St. Peter's Basilica.
Still Rome remained one of the strongholds of paganism, led by the aristocrats and senators. However, the new walls did not stop the city being sacked first by Alaric on 24 August 410, by Geiseric on 2 June 455, and even by general Ricimer's unpaid Roman troops (largely composed of barbarians) on 11 July 472. This was the first time in almost 800 years that Rome had fallen to an enemy. The previous sack of Rome had been accomplished by the Gauls under their leader Brennus in 387 BC. The sacking of 410 is seen as a major landmark in the decline and fall of the Western Roman Empire. St. Jerome, living in Bethlehem at the time, wrote that "The City which had taken the whole world was itself taken." These sackings of the city astonished all the Roman world. In any case, the damage caused by the sackings may have been overestimated. The population already started to decline from the late 4th century onward, although around the middle of the fifth century it seems that Rome continued to be the most populous city of the two parts of the Empire, with a population of no fewer than 650,000 inhabitants. The decline greatly accelerated following the capture of Africa Proconsularis by the Vandals. Many inhabitants now fled as the city no longer could be supplied with grain from Africa from the mid-5th century onward.
At the end of the 6th century Rome's population had reduced to around 30,000. Many monuments were being destroyed by the citizens themselves, who stripped stones from closed temples and other precious buildings, and even burned statues to make lime for their personal use. In addition, most of the increasing number of churches were built in this way. For example, the first Saint Peter's Basilica was erected using spoils from the abandoned Circus of Nero. This architectural cannibalism was a constant feature of Roman life until the Renaissance. From the 4th century, imperial edicts against stripping of stones and especially marble were common, but the need for their repetition shows that they were ineffective. Sometimes new churches were created by simply taking advantage of early Pagan temples, while sometimes changing the Pagan god or hero to a corresponding Christian saint or martyr. In this way, the Temple of Romulus and Remus became the basilica of the twin saints Cosmas and Damian. Later, the Pantheon, Temple of All Gods, became the church of All Martyrs.
Porta San Paolo, a gate in the Aurelian Walls, built between AD 271 and AD 275. During the Gothic Wars of the mid-6th century, Rome was besieged several times by Eastern Roman and Ostrogoth armies. Ostrogoths of Totila entered through this gate in 549, because of the treason of the Isaurian garrison.
In 480, the last Western Roman emperor, Julius Nepos, was murdered and a Roman general of barbarian origin, Odoacer, declared allegiance to Eastern Roman emperor Zeno. Despite owing nominal allegiance to Constantinople, Odoacer and later the Ostrogoths continued, like the last emperors, to rule Italy as a virtually independent realm from Ravenna. Meanwhile, the Senate, even though long since stripped of wider powers, continued to administer Rome itself, with the Pope usually coming from a senatorial family. This situation continued until Theodahad murdered Amalasuntha, a pro-imperial Gothic queen, and usurped the power in 535. The Eastern Roman emperor, Justinian I (reigned 527–565), used this as a pretext to send forces to Italy under his famed general Belisarius, recapturing the city next year, on 9 December AD 536. In 537–538, the Eastern Romans successfully defended the city in a year-long siege against the Ostrogothic army, and eventually took Ravenna, too.
Gothic resistance revived however, and on 17 December 546, the Ostrogoths under Totila recaptured and sacked Rome. Belisarius soon recovered the city, but the Ostrogoths retook it in 549. Belisarius was replaced by Narses, who captured Rome from the Ostrogoths for good in 552, ending the so-called Gothic Wars which had devastated much of Italy. The continual war around Rome in the 530s and 540s left it in a state of total disrepair – near-abandoned and desolate with much of its lower-lying parts turned into unhealthy marshes as the drainage systems were neglected and the Tiber's embankments fell into disrepair in the course of the latter half of the 6th century. Here, malaria developed. The aqueducts, except for one, were not repaired. The population, without imports of grain and oil from Sicily, shrank to less than 50,000 concentrated near the Tiber and around the Campus Martius, abandoning those districts without water supply. There is a legend, significant though untrue, that there was a moment where no one remained living in Rome.
Justinian I provided grants for the maintenance of public buildings, aqueducts and bridges—though, being mostly drawn from an Italy dramatically impoverished by the recent wars, these were not always sufficient. He also styled himself the patron of its remaining scholars, orators, physicians and lawyers in the stated hope that eventually more youths would seek a better education. After the wars, the Senate was theoretically restored, but under the supervision of the urban prefect and other officials appointed by, and responsible to, the Eastern Roman authorities in Ravenna.
However, the Pope was now one of the leading religious figures in the entire Byzantine Roman Empire and effectively more powerful locally than either the remaining senators or local Eastern Roman (Byzantine) officials. In practice, local power in Rome devolved to the Pope and, over the next few decades, both much of the remaining possessions of the senatorial aristocracy and the local Byzantine Roman administration in Rome were absorbed by the Church.
The reign of Justinian's nephew and successor Justin II (reigned 565–578) was marked from the Italian point of view by the invasion of the Lombards under Alboin (568). In capturing the regions of Benevento, Lombardy, Piedmont, Spoleto and Tuscany, the invaders effectively restricted Imperial authority to small islands of land surrounding a number of coastal cities, including Ravenna, Naples, Rome and the area of the future Venice. The one inland city continuing under Eastern Roman control was Perugia, which provided a repeatedly threatened overland link between Rome and Ravenna. In 578 and again in 580, the Senate, in some of its last recorded acts, had to ask for the support of Tiberius II Constantine (reigned 578–582) against the approaching Dukes, Faroald I of Spoleto and Zotto of Benevento.
Maurice (reigned 582–602) added a new factor in the continuing conflict by creating an alliance with Childebert II of Austrasia (reigned 575–595). The armies of the Frankish King invaded the Lombard territories in 584, 585, 588 and 590. Rome had suffered badly from a disastrous flood of the Tiber in 589, followed by a plague in 590. The latter is notable for the legend of the angel seen, while the newly elected Pope Gregory I (term 590–604) was passing in procession by Hadrian's Tomb, to hover over the building and to sheathe his flaming sword as a sign that the pestilence was about to cease. The city was safe from capture at least.
Agilulf, however, the new Lombard King (reigned 591 to c. 616), managed to secure peace with Childebert, reorganised his territories and resumed activities against both Naples and Rome by 592. With the Emperor preoccupied with wars in the eastern borders and the various succeeding Exarchs unable to secure Rome from invasion, Gregory took personal initiative in starting negotiations for a peace treaty. This was completed in the autumn of 598—later recognised by Maurice—lasting until the end of his reign.
The position of the Bishop of Rome was further strengthened under the usurper Phocas (reigned 602–610). Phocas recognised his primacy over that of the Patriarch of Constantinople and even decreed Pope Boniface III (607) to be "the head of all the Churches". Phocas's reign saw the erection of the last imperial monument in the Roman Forum, the column bearing his name. He also gave the Pope the Pantheon, at the time closed for centuries, and thus probably saved it from destruction.
During the 7th century, an influx of both Byzantine Roman officials and churchmen from elsewhere in the empire made both the local lay aristocracy and Church leadership largely Greek speaking. The population of Rome, a magnet for pilgrims, may have increased to 90,000. Eleven of thirteen popes between 678 and 752 were of Greek or Syrian descent. However, the strong Byzantine Roman cultural influence did not always lead to political harmony between Rome and Constantinople. In the controversy over Monothelitism, popes found themselves under severe pressure (sometimes amounting to physical force) when they failed to keep in step with Constantinople's shifting theological positions. In 653, Pope Martin I was deported to Constantinople and, after a show trial, exiled to the Crimea, where he died.
Then, in 663, Rome had its first imperial visit for two centuries, by Constans II—its worst disaster since the Gothic Wars when the Emperor proceeded to strip Rome of metal, including that from buildings and statues, to provide armament materials for use against the Saracens. However, for the next half century, despite further tensions, Rome and the Papacy continued to prefer continued Byzantine Roman rule: in part because the alternative was Lombard rule, and in part because Rome's food was largely coming from Papal estates elsewhere in the Empire, particularly Sicily.
Rome Timeline
772The Lombards briefly conquer Rome but Charlemagne liberates the city a year later.
800Charlemagne is crowned Holy Roman Emperor in St. Peter's Basilica.
846The Saracens sack St. Peter.
852Building of the Leonine Walls.
962Otto I crowned Emperor by Pope John XII
1000Emperor Otto III and Pope Sylvester II.
1084The Normans sack Rome.
1144Creation of the commune of Rome.
1300First Jubilee proclaimed by Pope Boniface VIII.
1303Foundation of the Roman University.
1309Pope Clement V moves the Holy Seat to Avignon.
1347Cola di Rienzo proclaims himself tribune.
1377Pope Gregory XI moves the Holy Seat back to Rome.
Break with Constantinople and formation of the Papal States
Further information: Duchy of Rome and Papal States
In 727, Pope Gregory II refused to accept the decrees of Emperor Leo III, which promoted the Emperor's iconoclasm. Leo reacted first by trying in vain to abduct the Pontiff, and then by sending a force of Ravennate troops under the command of the Exarch Paulus, but they were pushed back by the Lombards of Tuscia and Benevento. Byzantine general Eutychius sent west by the Emperor successfully captured Rome and restored it as a part of the empire in 728.
On 1 November 731, a council was called in St. Peter's by Gregory III to excommunicate the iconoclasts. The Emperor responded by confiscating large Papal estates in Sicily and Calabria and transferring areas previously ecclesiastically under the Pope to the Patriarch of Constantinople. Despite the tensions Gregory III never discontinued his support to the imperial efforts against external threats.
In this period the Lombard kingdom revived under the leadership of King Liutprand. In 730, he razed the countryside of Rome to punish the Pope, who had supported Duke Transamund II of Spoleto. Though still protected by his massive walls, the Pope could do little against the Lombard king, who managed to ally himself with the Byzantines. Other protectors were now needed. Gregory III was the first Pope to ask for concrete help from the Frankish Kingdom, then under the command of Charles Martel (739).
Liutprand's successor Aistulf was even more aggressive. He conquered Ferrara and Ravenna, ending the Exarchate of Ravenna. Rome seemed his next victim. In 754, Pope Stephen II went to France to name Pippin the Younger, king of the Franks, as patricius Romanorum, i.e. protector of Rome. In the August of that year the King and Pope together crossed back the Alps and defeated Aistulf at Pavia. When Pippin went back to St. Denis however, Aistulf did not keep his promises, and in 756 besieged Rome for 56 days. The Lombards returned north when they heard news of Pippin again moving to Italy. This time he agreed to give the Pope the promised territories, and the Papal States were born.
In 771 the new King of the Lombards, Desiderius, devised a plot to conquer Rome and seize Pope Stephen III during a feigned pilgrimage within its walls. His main ally was one Paulus Afiarta, chief of the Lombard party within the city. He conquered Rome in 772 but angered Charlemagne. However the plan failed, and Stephen's successor, Pope Hadrian I called Charlemagne against Desiderius, who was finally defeated in 773. The Lombard Kingdom was no more, and now Rome entered into the orbit of a new, greater political institution.
Numerous remains from this period, along with a museum devoted to Medieval Rome, can be seen at Crypta Balbi in Rome.
On 25 April 799 the new Pope, Leo III, led the traditional procession from the Lateran to the Church of San Lorenzo in Lucina along the Via Flaminia (now Via del Corso). Two nobles (followers of his predecessor Hadrian) who disliked the weakness of the Pope with regards to Charlemagne, attacked the processional train and delivered a life-threatening wound to the Pope. Leo fled to the King of the Franks, and in November, 800, the King entered Rome with a strong army and a number of French bishops. He declared a judicial trial to decide if Leo III were to remain Pope, or if the deposers' claims had reasons to be upheld. This trial, however, was only a part of a well thought out chain of events which ultimately surprised the world. The Pope was declared legitimate and the attempters subsequently exiled. On 25 December 800, Pope Leo III crowned Charlemagne Holy Roman Emperor in St. Peter's Basilica.
This act forever severed the loyalty of Rome from its imperial progenitor, Constantinople. It created instead a rival empire which, after a long series of conquests by Charlemagne, now encompassed most of the Christian Western territories.
Following the death of Charlemagne, the lack of a figure with equal prestige led the new institution into disagreement. At the same time the universal church of Rome had to face emergence of the lay interests of the City itself, spurred on by the conviction that the Roman people, though impoverished and abased, had again the right to elect the Western Emperor. The famous counterfeit document called the Donation of Constantine, prepared by the Papal notaries, guaranteed to the Pope a dominion stretching from Ravenna to Gaeta. This nominally included the suzerainty over Rome, but this was often highly disputed, and as the centuries passed, only the strongest Popes were to be able to assert it. The main element of weakness of the Papacy within the walls of the city was the continued necessity of the election of new popes, in which the emerging noble families soon managed to insert a leading role for themselves. The neighbouring powers, namely the Duchy of Spoleto and Toscana, and later the Emperors, learned how to take their own advantage of this internal weakness, playing the role of arbiters among the contestants.
Rome was indeed prey of anarchy in this age. The lowest point was touched in 897, when a raging crowd exhumed the corpse of a dead pope, Formosus, and put it on trial.
From 1048 to 1257, the papacy experienced increasing conflict with the leaders and churches of the Holy Roman Empire and the Byzantine (Eastern Roman) Empire. The latter culminated in the East-West Schism, dividing the Catholic Church and Eastern Orthodox Church. From 1257 to 1377, the pope, though the bishop of Rome, resided in Viterbo, Orvieto, and Perugia, and then Avignon. The return of the popes to Rome after the Avignon Papacy was followed by the Western Schism: the division of the western church between two, and for a time three, competing papal claimants.
In this period the renovated Church was again attracting pilgrims and prelates from all the Christian world, and money with them: even with a population of only 30,000, Rome was again becoming a city of consumers dependent upon the presence of a governmental bureaucracy. In the meantime, Italian cities were acquiring increasing autonomy, mainly led by new families which were replacing the old aristocracy with a new class formed by entrepreneurs, traders and merchants. After the sack of Rome by the Normans in 1084, the rebuilding of the city was supported by powerful families such as the Frangipane family and the Pierleoni family, whose wealth came from commerce and banking rather than landholdings. Inspired by neighbouring cities like Tivoli and Viterbo, Rome's people began to consider adopting a communal status and gaining a substantial amount of freedom from papal authority.
Led by Giordano Pierleoni, the Romans rebelled against the aristocracy and Church rule in 1143. The Senate and the Roman Republic, the Commune of Rome, were born again. Through the inflammatory words of preacher Arnaldo da Brescia, an idealistic, fierce opponent of ecclesiastical property and church interference in temporal affairs, the revolt that led to the creation of the Commune of Rome continued until it was put down in 1155, though it left its mark on the civil government of the Eternal City for centuries. 12th-century Rome, however, had little in common with the empire which had ruled over the Mediterranean some 700 years before, and soon the new Senate had to work hard to survive, choosing an ambiguous policy of shifting its support from the Pope to the Holy Roman Empire and vice versa as the political situation required. At Monteporzio, in 1167, during one of these shifts, in the war with Tusculum, Roman troops were defeated by the imperial forces of Frederick Barbarossa. Luckily, the winning enemies were soon dispersed by a plague and Rome was saved.
In 1188 the new communal government was finally recognised by Pope Clement III. The Pope had to make large cash payments to the communal officials, while the 56 senators became papal vassals. The Senate always had problems in the accomplishment of its function, and various changes were tried. Often a single Senator was in charge. This sometimes led to tyrannies, which did not help the stability of the newborn organism.
In 1204 the streets of Rome were again in flames when the struggle between Pope Innocent III's family and its rivals, the powerful Orsini family, led to riots in the city. Many ancient buildings were then destroyed by machines used by the rival bands to besiege their enemies in the innumerable towers and strongholds which were a hallmark of the Middle Age Italian towns.
The struggle between the Popes and the emperor Frederick II, also king of Naples and Sicily, saw Rome support the Ghibellines. To repay his loyalty, Frederick sent to the commune the Carroccio he had won from the Lombards at the battle of Cortenuova in 1234, and which was exposed in the Campidoglio.
In that year, during another revolt against the Pope, the Romans headed by senator Luca Savelli sacked the Lateran. Savelli was the father of Honorius IV, but in that age family ties often did not determine one's allegiance.
Rome was never to evolve into an autonomous, stable reign, as happened to other communes like Florence, Siena or Milan. The endless struggles between noble families (Savelli, Orsini, Colonna, Annibaldi), the ambiguous position of the Popes, the haughtiness of a population which never abandoned the dreams of their splendid past but, at the same time, thought only of immediate advantage, and the weakness of the republican institutions always deprived the city of this possibility.
In an attempt to imitate more successful communes, in 1252 the people elected a foreign Senator, the Bolognese Brancaleone degli Andalò. In order to bring peace in the city he suppressed the most powerful nobles (destroying some 140 towers), reorganised the working classes and issued a code of laws inspired by those of northern Italy. Brancaleone was a tough figure, but died in 1258 with almost nothing of his reforms turned into reality. Five years later Charles I of Anjou, then king of Naples, was elected Senator. He entered the city only in 1265, but soon his presence was needed to face Conradin, the Hohenstaufen's heir who was coming to claim his family's rights over southern Italy, and left the city. After June 1265 Rome was again a democratic republic, electing Henry of Castile as senator. But Conradin and the Ghibelline party were crushed in the Battle of Tagliacozzo (1268), and therefore Rome fell again in the hands of Charles.
Nicholas III, a member of Orsini family, was elected in 1277 and moved the seat of the Popes from the Lateran to the more defensible Vatican. He also ordered that no foreigner could become senator of Rome. Being a Roman himself, he had himself elected senator by the people. With this move, the city began again to side for the papal party. In 1285 Charles was again Senator, but the Sicilian Vespers reduced his charisma, and the city was thenceforth free from his authority. The next senator was again a Roman, and again a pope, Honorius IV of the Savelli.
The successor to Celestine V was a Roman of the Caetani family, Boniface VIII. Entangled in a local feud against the traditional rivals of his family, the Colonna, at the same time he struggled to assure the universal supremacy of the Holy See. In 1300 he launched the first Jubilee and in 1303 founded the first University of Rome. The Jubilee was an important move for Rome, as it further increased its international prestige and, most of all, the city's economy was boosted by the flow of pilgrims. Boniface died in 1303 after the humiliation of the Schiaffo di Anagni ("Slap of Anagni"), which signalled instead the rule of the King of France over the Papacy and marked another period of decline for Rome.
Boniface's successor, Clement V, never entered the city, starting the so-called "Avignon captivity", the absence of the Popes from their Roman seat in favour of Avignon, which would last for more than 70 years. This situation brought the independence of the local powers, but these were revealed to be largely unstable; and the lack of the holy revenues caused a deep decay of Rome. For more than a century Rome had no new major buildings. Furthermore, many of the monuments of the city, including the main churches, began to fall into ruin.
Cola di Rienzo stormed the Capitoline Hill in 1347 to create a new Roman Republic. Though short-lived, his attempt is recorded by a 19th-century statue near the ramped Cordonata leading to Michelangelo's Piazza del Campidoglio.
In spite of its decline and the absence of the Pope, Rome had not lost its spiritual prestige: in 1341 the famous poet Petrarca came to the city to be crowned as Poet laureate in Capitoline Hill. Noblemen and poor people at one time demanded with one voice the return of the Pope. Among the many ambassadors that in this period took their way to Avignon, emerged the bizarre but eloquent figure of Cola di Rienzo. As his personal power among the people increased by time, on 20 May 1347 he conquered the Capitoline at the head of an enthusiastic crowd. The period of his power, though very short-lived, aspired to the prestige of Ancient Rome. Now in possession of dictatorial powers, he took the title of "tribune", referring to the pleb's magistracy of the Roman Republic. Cola also considered himself at an equal status of that of the Holy Roman Emperor. On 1 August, he conferred Roman citizenship on all the Italian cities, and even prepared for the election of a Roman emperor of Italy. It was too much: the Pope denounced him as heretic, criminal and pagan, the populace had begun to be disenchanted with him, while the nobles had always hated him. On 15 December, he was forced to flee.
In August 1354, Cola was again a protagonist, when Cardinal Gil Alvarez De Albornoz entrusted him with the role of "senator of Rome" in his programme of reassuring the Pope's rule in the Papal States. In October the tyrannical Cola, who had become again very unpopular for his delirious behaviour and heavy bills, was killed in a riot provoked by the powerful family of the Colonna. In April 1355, Charles IV of Bohemia entered the city for the ritual coronation as Emperor. His visit was very disappointing for the citizens. He had little money, received the crown not from the Pope but from a Cardinal, and moved away after a few days.
With the emperor back in his lands, Albornoz could regain a certain control over the city, while remaining in his safe citadel in Montefiascone, in the Northern Lazio. The senators were chosen directly by the Pope from several cities of Italy, but the city was in fact independent. The Senate council included six judges, five notaries, six marshals, several familiars, twenty knights and twenty armed men. Albornoz had heavily suppressed the traditional aristocratic families, and the "democratic" party felt confident enough to start an aggressive policy. In 1362 Rome declared war on Velletri. This move, however, provoked a civil war. The countryside party hired a condottieri band called "Del Cappello" ("Hat"), while the Romans bought the services of German and Hungarian troops, plus a citizen levy of 600 knights and even 22,000 infantry. This was the period in which condottieri bands were active in Italy. Many of the Savelli, Orsini and Annibaldi expelled from Rome became leaders of such military units. The war with Velletri languished, and Rome again gave itself to the new Pope, Urban V, provided Albornoz did not enter the walls.
On 16 October 1367, in reply to the prayers of St Brigid and Petrarca, Urban finally visited for the city. During his presence, Charles IV was again crowned in the city (October 1368). In addition, the Byzantine emperor John V Palaeologus came in Rome to beg for a crusade against the Ottoman Empire, but in vain. However, Urban did not like the unhealthy air of the city, and on 5 September 1370 he sailed again to Avignon. His successor, Gregory XI, officially set the date of his return to Rome at May 1372, but again the French cardinals and the King stopped him.
Only on 17 January 1377, Gregory XI could finally reinstate the Holy See in Rome.
The incoherent behaviour of his successor, the Italian Urban VI, provoked in 1378 the Western Schism, which impeded any true attempt of improving the conditions of the decaying Rome. The 14th century, with the absence of the popes during the Avignon Papacy, had been a century of neglect and misery for the city of Rome, which dropped to its lowest level of population. With the return of the papacy to Rome repeatedly postponed because of the bad conditions of the city and the lack of control and security, it was first necessary to strengthen the political and doctrinal aspects of the pontiff.
When in 1377 Gregory XI was in fact returned to Rome, he found a city in anarchy because of the struggles between the nobility and the popular faction, and in which his power was now more formal than real. There followed four decades of instability, characterised by the local power struggle between the commune and the papacy, and internationally by the great Western Schism, at the end of which was elected Pope, Martin V. He restored order, laying the foundations of its rebirth.
In 1433 the Duke of Milan, Filippo Maria Visconti signed a peace treaty with Florence and Venice. He then sent the condottieri Niccolò Fortebraccio and Francesco Sforza to harass the Papal States, in vengeance for Eugene IV's support to the two former republics.
Fortebraccio, supported by the Colonna, occupied Tivoli in October 1433 and ravaged Rome's countryside. Despite the concessions made by Eugene to the Visconti, the Milanese soldiers did not stop their destruction. This led the Romans, on 29 May 1434 to institute a Republican government under the Banderesi. Eugene left the city a few days later, during the night of 4 June.
However, the Banderesi proved incapable of governing the city, and their inadequacies and violence soon deprived them of popular support. The city was therefore returned to Eugene by the army of Giovanni Vitelleschi on 26 October 1434. After the death in mysterious circumstances of Vitelleschi, the city came under the control of Ludovico Scarampo, Patriarch of Aquileia. Eugene returned to Rome on 28 September 1443.
The latter half of the 15th century saw the seat of the Italian Renaissance move to Rome from Florence. The Papacy wanted to surpass the grandeur of other Italian cities. To this end the popes created increasingly extravagant churches, bridges, town squares and public spaces, including a new Saint Peter's Basilica, the Sistine Chapel, Ponte Sisto (the first bridge to be built across the Tiber since antiquity), and Piazza Navona. The Popes were also patrons of the arts engaging such artists as Michelangelo, Perugino, Raphael, Ghirlandaio, Luca Signorelli, Botticelli, and Cosimo Rosselli.
Under Pope Nicholas V, who became Pontiff on 19 March 1447, the Renaissance can be said to have begun in Rome, heralding a period in which the city became the centre of Humanism. He was the first Pope to embellish the Roman court with scholars and artists, including Lorenzo Valla and Vespasiano da Bisticci.
On 4 September 1449 Nicholas proclaimed a Jubilee for the following year, which saw a great influx of pilgrims from all Europe. The crowd was so large that in December, on Ponte Sant'Angelo, some 200 people died, crushed underfoot or drowned in the River Tiber. Later that year the Plague reappeared in the city, and Nicholas fled.
However Nicholas brought stability to the temporal power of the Papacy, a power in which the Emperor was to have no part at all. In this way, the coronation and the marriage of Frederick III, Holy Roman Emperor on 16 March 1452, was more a civil ceremony. The Papacy now controlled Rome with a strong hand. A plot by Stefano Porcari, whose aim was the restoration of the Republic, was ruthlessly suppressed in January 1453. Porcari was hanged together with the other plotters, Francesco Gabadeo, Pietro de Monterotondo, Battista Sciarra and Angiolo Ronconi, but the Pope gained a treacherous reputation, as when the execution was beginning he was too drunk to confirm the grace he had previously given to Sciarra and Ronconi.
Nicholas was also actively involved in Rome's urban renewal, in collaboration with Leon Battista Alberti, including the construction of a new St Peter's Basilica.
Nicholas' successor Calixtus III neglected Nicholas's cultural policies, instead devoting himself to his greatest passion, his nephews. The Tuscan Pius II, who took the reins after his death in 1458, was a great Humanist, but did little for Rome. During his reign Lorenzo Valla demonstrated that the Donation of Constantine was a forgery. Pius was the first Pope to use guns, in campaign against the rebel barons Savelli in the neighbourhood of Rome, in 1461. One year later the bringing to Rome of the head of the Apostle St. Andrew produced a great number of pilgrims. The reign of Pope Paul II (1464–1471) was notable only for the reintroduction of the Carnival, which was to become a very popular feast in Rome in the following centuries. In the same year (1468) a plot against the Pope was uncovered, organised by the intellectuals of the Roman Academy founded by Pomponio Leto. The conspirators were sent to Castel Sant'Angelo.
More important by far was the Pontificate of Sixtus IV, considered the first Pope-King of Rome. In order to favour his relative Girolamo Riario, he promoted the unsuccessful Congiura dei Pazzi against the Medici of Florence (26 April 1478) and in Rome fought the Colonna and the Orsini. The personal politics of intrigue and war required much money, but in spite of this Sixtus was a true patron of art in the manner of Nicholas V. He reopened the Academy and reorganised the Collegio degli Abbreviatori, and in 1471 began the construction of the Vatican Library, whose first curator was Platina. The Library was officially founded on 15 Jun
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Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.
The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.
The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.
Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.
There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.
Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.
Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.
Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.
Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.
Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.
All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.
Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.
After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.
Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.
Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).
Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.
Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.
Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.
Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).
Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.
So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).
Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.
The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.
Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.
In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.
Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.
Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.
Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.
The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.
The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.
The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.
The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.
The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.
Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.
Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.
Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.
The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.
The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.
Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.
Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.
Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.
The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.
Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.
Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.
Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.
The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.
The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.
The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.
The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).
The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.
Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.
There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.
Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.
Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.
As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.
The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).
The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.
Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.
Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.
Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.
Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.
A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.
An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.
MTA Chair & CEO Janno Lieber announces the release of Reduced Fare OMNY cards on Friday, Dec 13, 2024 at the Stone Street Customer Service Center.
(Marc A. Hermann / MTA)
Reducing Food Loss and Waste: Advancing the Agenda
September 20, 2017 | New York, NY
Pictured (L-R):
* Maria Fernanda Mejia, President, Kellogg Latin America
* Lorna Donatone, CEO, Geographic Regions and Region Chair for North America, Sodexo
This Champions 12.3-hosted event assessed global progress toward Target 12.3, announced landmark developments and set forth a road to success. The event coincided with the 72nd UN General Assembly and Climate Week.
Find out more at www.wri.org/events/2017/09/reducing-food-loss-and-waste-a...
Photo credit: Robert Carlo | New York for media, publication, and digital outsources. @robertcarlonewyork for all Social Media
Top layer of black lacy net; underdress of reddish black silk. It rustles. It's nice. And it was reduced from £120 to £18 (Downtown, Grantham). Oh joy...
During its annual retreat, the Delaware Cancer Consortium and physical educator teachers from across the state heard from Gov. Carney, Lt. Gov. Hall-Long, Secretary Walker and national experts on the role physical activity plays in reducing the risk of cancer. Creating a culture of prevention through physical activity, particularly among the young, was the theme of the April 25 retreat.
Dr. I-Min Lee, a professor in the Department of Epidemiology at Harvard, said only 27% of high school students and 20% of adults in Delaware get the daily recommended amount of physical activity. Dr. Lee said that the habit of exercise “follows you when you become an adult” and helps to reduce your risk for the vast number of cancers. “Prevention is hard,” Dr. Lee said, “and it’s hard to show the difference it makes.”
Secretary Walker said a 2014 Nemours survey of 2,600 Delaware households with children found that 36% of children ages 2 to 17 were overweight or obese. “We as a state can do better,” she said. And that 54% of children ages 2 to 17 consumed 5 or more servings of fruits and veggies per day. “Again, with good health promotion, we can push that number even higher,” Secretary Walker said.
Progress was made in the reduction of children consuming 2 cups of sugar-sweetened beverages per week and the percentage of Delaware babies who were ever breastfed.
The retreat’s keynote address was delivered by Tom Farey, executive director of the Sports and Society Program at the Aspen Institute. Farey’s work has found that physical inactivity and obesity are risk factors for cancer and other ailments.
To learn more about how you can reduce your risk of cancer: