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Hydra (genus)

Hydra is a genus of small freshwater hydrozoans of the phylum Cnidaria. They are native to the temperate and tropical regions. The genus was named by Linnaeus in 1758 after the Hydra, which was the many-headed beast of myth defeated by Heracles, as when the animal has a part severed, it will regenerate much like the mythical hydra’s heads. Biologists are especially interested in Hydra because of their regenerative ability; they do not appear to die of old age, or to age at all.

Morphology

Hydra has a tubular, radially symmetric body up to 10 mm long when extended, secured by a simple adhesive foot known as the basal disc. Gland cells in the basal disc secrete a sticky fluid that accounts for its adhesive properties.

At the free end of the body is a mouth opening surrounded by one to twelve thin, mobile tentacles. Each tentacle, or cnida (plural: cnidae), is clothed with highly specialised stinging cells called cnidocytes. Cnidocytes contain specialized structures called nematocysts, which look like miniature light bulbs with a coiled thread inside. At the narrow outer edge of the cnidocyte is a short trigger hair called a cnidocil. Upon contact with prey, the contents of the nematocyst are explosively discharged, firing a dart-like thread containing neurotoxins into whatever triggered the release. This can paralyze the prey, especially if many hundreds of nematocysts are fired.

Hydra has two main body layers, which makes it "diploblastic". The layers are separated by mesoglea, a gel-like substance. The outer layer is the epidermis, and the inner layer is called the gastrodermis, because it lines the stomach. The cells making up these two body layers are relatively simple. Hydramacin is a bactericide recently discovered in Hydra; it protects the outer layer against infection. A single Hydra is composed of 50,000 to 100,000 cells which consist of three specific stem cell populations that create many different cell types. These stem cells continually renew themselves in the body column. Hydras have two significant structures on their body: the "head" and the "foot". When a Hydra is cut in half, each half regenerates and forms into a small Hydra; the "head" regenerates a "foot" and the "foot" regenerates a "head". If the Hydra is sliced into many segments then the middle slices form both a "head" and a "foot".

Respiration and excretion occur by diffusion throughout the surface of the epidermis, while larger excreta are discharged through the mouth.

Nervous system
The nervous system of Hydra is a nerve net, which is structurally simple compared to more derived animal nervous systems. Hydra does not have a recognizable brain or true muscles. Nerve nets connect sensory photoreceptors and touch-sensitive nerve cells located in the body wall and tentacles.

The structure of the nerve net has two levels:
 * level 1 – sensory cells or internal cells; and
 * level 2 – interconnected ganglion cells synapsed to epithelial or motor cells.

Some have only two sheets of neurons.

Motion and locomotion
If Hydra are alarmed or attacked, the tentacles can be retracted to small buds, and the body column itself can be retracted to a small gelatinous sphere. Hydra generally react in the same way regardless of the direction of the stimulus, and this may be due to the simplicity of the nerve nets.

Hydra are generally sedentary or sessile, but do occasionally move quite readily, especially when hunting. They have two distinct methods for moving – 'looping' and 'somersaulting'. They do this by bending over and attaching themselves to the substrate with the mouth and tentacles and then relocate the foot, which provides the usual attachment, this process is called looping. In somersaulting, the body then bends over and makes a new place of attachment with the foot. By this process of "looping" or "somersaulting", a Hydra can move several inches (c. 100 mm) in a day. Hydra may also move by amoeboid motion of their bases or by detaching from the substrate and floating away in the current.

Reproduction and life cycle
Most hydra species do not have any gender system. Instead, when food is plentiful, many Hydra reproduce asexually by budding. The buds form from the body wall, grow into miniature adults and break away when mature.

When a hydra is well fed, a new bud can form every two days. When conditions are harsh, often before winter or in poor feeding conditions, sexual reproduction occurs in some Hydra. Swellings in the body wall develop into either ovaries or testes. The testes release free-swimming gametes into the water, and these can fertilize the egg in the ovary of another individual. The fertilized eggs secrete a tough outer coating, and, as the adult dies (due to starvation or cold), these resting eggs fall to the bottom of the lake or pond to await better conditions, whereupon they hatch into nymph Hydra. Some Hydra species, like Hydra circumcincta and Hydra viridissima, are hermaphrodites and may produce both testes and ovaries at the same time.

Many members of the Hydrozoa go through a body change from a polyp to an adult form called a medusa, which is usually the life stage where sexual reproduction occurs, but Hydra do not progress beyond the polyp phase.

Feeding
Hydra mainly feed on aquatic invertebrates such as Daphnia and Cyclops.

While feeding, Hydra extend their body to maximum length and then slowly extend their tentacles. Despite their simple construction, the tentacles of Hydra are extraordinarily extensible and can be four to five times the length of the body. Once fully extended, the tentacles are slowly maneuvered around waiting for contact with a suitable prey animal. Upon contact, nematocysts on the tentacle fire into the prey, and the tentacle itself coils around the prey. Most of the tentacles join in the attack within 30 seconds to subdue the struggling prey. Within two minutes, the tentacles surround the prey and move it into the open mouth aperture. Within ten minutes, the prey is engulfed within the body cavity, and digestion commences. Hydra can stretch their body wall considerably.

The feeding behaviour of Hydra demonstrates the sophistication of what appears to be a simple nervous system.

Some species of Hydra exist in a mutual relationship with various types of unicellular algae. The algae are protected from predators by Hydra; in return, photosynthetic products from the algae are beneficial as a food source to Hydra , and even help to maintain the Hydra microbiome.

Measuring the feeding response
The feeding response in Hydra is induced by glutathione (specifically in the reduced state as GSH) released from damaged tissue of injured prey. There are several methods conventionally used for quantification of the feeding response. In some, the duration for which the mouth remains open is measured. Other methods rely on counting the number of Hydra among a small population showing the feeding response after addition of glutathione. Recently, an assay for measuring the feeding response in hydra has been developed. In this method, the linear two-dimensional distance between the tip of the tentacle and the mouth of hydra was shown to be a direct measure of the extent of the feeding response. This method has been validated using a starvation model, as starvation is known to cause enhancement of the Hydra feeding response.

Predators
The species Hydra oligactis is preyed upon by the flatworm Microstomum lineare.

Tissue regeneration
Hydras undergo morphallaxis (tissue regeneration) when injured or severed. Typically, Hydras reproduce by just budding off a whole new individual; the bud occurs around two-thirds of the way down the body axis. When a Hydra is cut in half, each half regenerates and forms into a small Hydra; the "head" regenerates a "foot" and the "foot" regenerates a "head". This regeneration occurs without cell division. If the Hydra is sliced into many segments, the middle slices form both a "head" and a "foot". The polarity of the regeneration is explained by two pairs of positional value gradients. There is both a head and foot activation and inhibition gradient. The head activation and inhibition works in an opposite direction of the pair of foot gradients. The evidence for these gradients was shown in the early 1900s with grafting experiments. The inhibitors for both gradients have shown to be important to block the bud formation. The location where the bud forms is where the gradients are low for both the head and foot.

Hydras are capable of regenerating from pieces of tissue from the body and additionally after tissue dissociation from reaggregates. This process takes place not only in the pieces of tissue excised from the body column, but also from re-aggregates of dissociated single cells. It was found that in these aggregates, cells initially distributed randomly undergo sorting and form two epithelial cell layers, in which the endodermal epithelial cells play more active roles in the process. Active mobility of these endodermal epithelial cells forms two layers in both the re-aggregate and the re-generating tip of the excised tissue. As these two layers are established, a patterning process takes place to form heads and feet.

Non-senescence
Daniel Martinez claimed in a 1998 article in Experimental Gerontology that Hydra are biologically immortal. This publication has been widely cited as evidence that Hydra do not senesce (do not age), and that they are proof of the existence of non-senescing organisms generally. In 2010, Preston Estep published (also in Experimental Gerontology) a letter to the editor arguing that the Martinez data refutes the hypothesis that Hydra do not senesce.

The controversial unlimited lifespan of Hydra has attracted much attention from scientists. Research today appears to confirm Martinez' study. Hydra stem cells have a capacity for indefinite self-renewal. The transcription factor "forkhead box O" (FoxO) has been identified as a critical driver of the continuous self-renewal of Hydra. In experiments, a drastically reduced population growth resulted from FoxO down-regulation.

In bilaterally symmetrical organisms (Bilateria), the transcription factor FoxO affects stress response, lifespan, and increase in stem cells. If this transcription factor is knocked down in bilaterian model organisms, such as fruit flies and nematodes, their lifespan is significantly decreased. In experiments on H. vulgaris (a radially symmetrical member of phylum Cnidaria), when FoxO levels were decreased, there was a negative effect on many key features of the Hydra, but no death was observed, thus it is believed other factors may contribute to the apparent lack of aging in these creatures.

DNA repair
Hydra are capable of two types of DNA repair: nucleotide excision repair and base excision repair. The repair pathways facilitate DNA replication by removing DNA damage. Their identification in hydra was based, in part, on the presence in its genome of genes homologous to ones present in other genetically well studied species playing key roles in these DNA repair pathways.

Genomics
An ortholog comparison analysis done within the last decade demonstrated that Hydra share a minimum of 6,071 genes with humans. Hydra is becoming an increasingly better model system as more genetic approaches become available. Transgenic hydra have become attractive model organisms to study the evolution of immunity. A draft of the genome of Hydra magnipapillata was reported in 2010.

The genomes of cnidarians are usually less than 500 Mb (megabases) in size, as in the Hydra viridissima, which has a genome size of approximately 300 Mb. In contrast, the genomes of brown hydras are approximately 1 Gb in size. This is because the brown hydra genome is the result of an expansion event involving LINEs, a type of transposable elements, in particular, a single family of the CR1 class. This expansion is unique to this subgroup of the genus Hydra and is absent in the green hydra, which has a repeating landscape similar to other cnidarians. These genome characteristics make Hydra attractive for studies of transposon-driven speciations and genome expansions.

Due to the simplicity of their life cycle when compared to other hydrozoans, hydras have lost many genes that correspond to cell types or metabolic pathways of which the ancestral function is still unknown.

Hydra genome shows a preference towards proximal promoters. Thanks to this feature, many reporter cell lines have been created with regions around 500 to 2000 bases upstream of the gene of interest. Its cis-regulatory elements (CRE) are mostly located less than 2000 base pairs upstream from the closest transcription initiation site, but there are CREs located further away.

Its chromatin has a Rabl configuration. There are interactions between the centromeres of different chromosomes and the centromeres and telomeres of the same chromosome. It presents a great number of intercentromeric interactions when compared to other cnidarians, probably due to the loss of multiple subunits of condensin II. It is organized in domains that span dozens to hundreds of megabases, containing epigenetically co-regulated genes and flanked by boundaries located within heterochromatin.

Transcriptomics
Different Hydra cell types express gene families of different evolutionary ages. Progenitor cells (stem cells, neuron and nematocyst precursors, and germ cells) express genes from families that predate metazoans. Among differentiated cells some express genes from families that date from the base of metazoans, like gland and neuronal cells, and others express genes from newer families, originating from the base of cnidaria or medusozoa, like nematocysts. Interstitial cells contain translation factors with a function that has been conserved for at least 400 million years.

title: Hydra (genus)


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Hydrus

Hydrus is a small constellation in the deep southern sky. It was one of twelve constellations created by Petrus Plancius from the observations of Pieter Dirkszoon Keyser and Frederick de Houtman and it first appeared on a 35-cm (14 in) diameter celestial globe published in late 1597 (or early 1598) in Amsterdam by Plancius and Jodocus Hondius. The first depiction of this constellation in a celestial atlas was in Johann Bayer's Uranometria of 1603. The French explorer and astronomer Nicolas Louis de Lacaille charted the brighter stars and gave their Bayer designations in 1756. Its name means "male water snake", as opposed to Hydra, a much larger constellation that represents a female water snake. It remains below the horizon for most Northern Hemisphere observers.

The brightest star is the 2.8-magnitude Beta Hydri, also the closest reasonably bright star to the south celestial pole. Pulsating between magnitude 3.26 and 3.33, Gamma Hydri is a variable red giant 60 times the diameter of the Sun. Lying near it is VW Hydri, one of the brightest dwarf novae in the heavens. Four star systems in Hydrus have been found to have exoplanets to date, including HD 10180, which could bear up to nine planetary companions.

History
Hydrus was one of the twelve constellations established by the astronomer Petrus Plancius from the observations of the southern sky by the Dutch explorers Pieter Dirkszoon Keyser and Frederick de Houtman, who had sailed on the first Dutch trading expedition, known as the Eerste Schipvaart, to the East Indies. It first appeared on a 35-cm (14 in) diameter celestial globe published in late 1597 (or early 1598) in Amsterdam by Plancius with Jodocus Hondius. The first depiction of this constellation in a celestial atlas was in the German cartographer Johann Bayer's Uranometria of 1603. De Houtman included it in his southern star catalogue the same year under the Dutch name De Waterslang, "The Water Snake", it representing a type of snake encountered on the expedition rather than a mythical creature. The French explorer and astronomer Nicolas Louis de Lacaille called it l’Hydre Mâle on the 1756 version of his planisphere of the southern skies, distinguishing it from the feminine Hydra. The French name was retained by Jean Fortin in 1776 for his Atlas Céleste, while Lacaille Latinised the name to Hydrus for his revised Coelum Australe Stelliferum in 1763.

Characteristics
Irregular in shape, Hydrus is bordered by Mensa to the southeast, Eridanus to the east, Horologium and Reticulum to the northeast, Phoenix to the north, Tucana to the northwest and west, and Octans to the south; Lacaille had shortened Hydrus' tail to make space for this last constellation he had drawn up. Covering 243 square degrees and 0.589% of the night sky, it ranks 61st of the 88 constellations in size. The three-letter abbreviation for the constellation, as adopted by the International Astronomical Union in 1922, is "Hyi". The official constellation boundaries, as set by Belgian astronomer Eugène Delporte in 1930, are defined by a polygon of 12 segments. In the equatorial coordinate system, the right ascension coordinates of these borders lie between 00:06.1:0 and 04:35.1:0, while the declination coordinates are between −57.85° and −82.06°. As one of the deep southern constellations, it remains below the horizon at latitudes north of the 30th parallel in the Northern Hemisphere, and is circumpolar at latitudes south of the 50th parallel in the Southern Hemisphere. Herman Melville mentions it and Argo Navis in Moby Dick "beneath effulgent Antarctic Skies", highlighting his knowledge of the southern constellations from whaling voyages. A line drawn between the long axis of the Southern Cross to Beta Hydri and then extended 4.5 times will mark a point due south. Hydrus culminates at midnight around 26 October.

Stars
Keyzer and de Houtman assigned fifteen stars to the constellation in their Malay and Madagascan vocabulary, with a star that would be later designated as Alpha Hydri marking the head, Gamma the chest and a number of stars that were later allocated to Tucana, Reticulum, Mensa and Horologium marking the body and tail. Lacaille charted and designated 20 stars with the Bayer designations Alpha through to Tau in 1756. Of these, he used the designations Eta, Pi and Tau twice each, for three sets of two stars close together, and omitted Omicron and Xi. He assigned Rho to a star that subsequent astronomers were unable to find.

Beta Hydri, the brightest star in Hydrus, is a yellow star of apparent magnitude 2.8, lying 24 light-years from Earth. It has about 104% of the mass of the Sun and 181% of the Sun's radius, with more than three times the Sun's luminosity. The spectrum of this star matches a stellar classification of G2 IV, with the luminosity class of 'IV' indicating this is a subgiant star. As such, it is a slightly more evolved star than the Sun, with the supply of hydrogen fuel at its core becoming exhausted. It is the nearest subgiant star to the Sun and one of the oldest stars in the solar neighbourhood. Thought to be between 6.4 and 7.1 billion years old, this star bears some resemblance to what the Sun may look like in the far distant future, making it an object of interest to astronomers. It is also the closest bright star to the south celestial pole.

Located at the northern edge of the constellation and just southwest of Achernar is Alpha Hydri, a white sub-giant star of magnitude 2.9, situated 72 light-years from Earth. Of spectral type F0IV, it is beginning to cool and enlarge as it uses up its supply of hydrogen. It is twice as massive and 3.3 times as wide as the Sun and 26 times more luminous. A line drawn between Alpha Hydri and Beta Centauri is bisected by the south celestial pole.

In the southeastern corner of the constellation is Gamma Hydri, a red giant of spectral type M2III located 214 light-years from Earth. It is a semi-regular variable star, pulsating between magnitudes 3.26 and 3.33. Observations over five years were not able to establish its periodicity. It is around 1.5 to 2 times as massive as the Sun, and has expanded to about 60 times the Sun's diameter. It shines with about 655 times the luminosity of the Sun. Located 3° northeast of Gamma is the VW Hydri, a dwarf nova of the SU Ursae Majoris type. It is a close binary system that consists of a white dwarf and another star, the former drawing off matter from the latter into a bright accretion disk. These systems are characterised by frequent eruptions and less frequent supereruptions. The former are smooth, while the latter exhibit short "superhumps" of heightened activity. One of the brightest dwarf novae in the sky, it has a baseline magnitude of 14.4 and can brighten to magnitude 8.4 during peak activity. BL Hydri is another close binary system composed of a low-mass star and a strongly magnetic white dwarf. Known as a polar or AM Herculis variable, these produce polarized optical and infrared emissions and intense soft and hard X-ray emissions to the frequency of the white dwarf's rotation period—in this case 113.6 minutes.

There are two notable optical double stars in Hydrus. Pi Hydri, composed of Pi1 Hydri and Pi2 Hydri, is divisible in binoculars. Around 476 light-years distant, Pi1 is a red giant of spectral type M1III that varies between magnitudes 5.52 and 5.58. Pi2 is an orange giant of spectral type K2III and shining with a magnitude of 5.7, around 488 light-years from Earth.

Eta Hydri is the other optical double, composed of Eta1 and Eta2. Eta1 is a blue-white main sequence star of spectral type B9V that was suspected of being variable, and is located just over 700 light-years away. Eta2 has a magnitude of 4.7 and is a yellow giant star of spectral type G8.5III around 218 light-years distant, which has evolved off the main sequence and is expanding and cooling on its way to becoming a red giant. Calculations of its mass indicate it was most likely a white A-type main sequence star for most of its existence, around twice the mass of the Sun. A planet, Eta2 Hydri b, greater than 6.5 times the mass of Jupiter was discovered in 2005, orbiting around Eta2 every 711 days at a distance of 1.93 astronomical units (AU).

Three other systems have been found to have planets, most notably the Sun-like star HD 10180, which has seven planets, plus possibly an additional two for a total of nine—as of 2012 more than any other system to date, including the Solar System. Lying around 127 ly from the Earth, it has an apparent magnitude of 7.33.

GJ 3021 is a solar twin—a star very like the Sun—around 57 light-years distant with a spectral type G8V and magnitude of 6.7. It has a Jovian planet companion (GJ 3021 b). Orbiting about 0.5 AU from its star, it has a minimum mass 3.37 times that of Jupiter and a period of around 133 days. The system is a complex one as the faint star GJ 3021B orbits at a distance of 68 AU; it is a red dwarf of spectral type M4V.

HD 20003 is a star of magnitude 8.37. It is a yellow main sequence star of spectral type G8V a little cooler and smaller than the Sun around 143 light-years away. It has two planets that are around 12 and 13.5 times as massive as the Earth with periods of just under 12 and 34 days respectively.

Deep-sky objects
Hydrus contains only faint deep-sky objects. IC 1717 was a deep-sky object discovered by the Danish astronomer John Louis Emil Dreyer in the late 19th century. The object at the coordinate Dreyer observed is no longer there, and is now a mystery. It was very likely to have been a faint comet. PGC 6240, known as the White Rose Galaxy, is a giant spiral galaxy surrounded by shells resembling rose petals, located around 345 million light years from the Solar System. Unusually, it has cohorts of globular clusters of three distinct ages suggesting bouts of post-starburst formation following a merger with another galaxy. The constellation also contains a spiral galaxy, NGC 1511, which lies edge on to observers on Earth and is readily viewed in amateur telescopes.

Located mostly in Dorado, the Large Magellanic Cloud extends into Hydrus. The globular cluster NGC 1466 is an outlying component of the galaxy, and contains many RR Lyrae-type variable stars. It has a magnitude of 11.59 and is thought to be over 12 billion years old. Two stars, HD 24188 of magnitude 6.3 and HD 24115 of magnitude 9.0, lie nearby in its foreground. NGC 602 is composed of an emission nebula and a young, bright open cluster of stars that is an outlying component on the eastern edge of the Small Magellanic Cloud, a satellite galaxy to the Milky Way. Most of the cloud is located in the neighbouring constellation Tucana.

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Hercules

Hercules is the Roman equivalent of the Greek divine hero Heracles, son of Jupiter and the mortal Alcmena. In classical mythology, Hercules is famous for his strength and for his numerous far-ranging adventures.

The Romans adapted the Greek hero's iconography and myths for their literature and art under the name Hercules. In later Western art and literature and in popular culture, Hercules is more commonly used than Heracles as the name of the hero. Hercules is a multifaceted figure with contradictory characteristics, which enabled later artists and writers to pick and choose how to represent him. This article provides an introduction to representations of Hercules in the later tradition.

Birth and early life
In Roman mythology, although Hercules was seen as the champion of the weak and a great protector, his personal problems started at birth. Juno sent two witches to prevent the birth, but they were tricked by one of Alcmene's servants and sent to another room. Juno then sent serpents to kill him in his cradle, but Hercules strangled them both. In one version of the myth, Alcmene abandoned her baby in the woods in order to protect him from Juno's wrath, but he was found by the goddess Minerva who brought him to Juno, claiming he was an orphan child left in the woods who needed nourishment. Juno suckled Hercules at her own breast until the infant bit her nipple, at which point she pushed him away, spilling her milk across the night sky and so forming the Milky Way. She then gave the infant back to Minerva and told her to take care of the baby herself. In feeding the child from her own breast, the goddess inadvertently imbued him with further strength and power.

Roman era

The Latin name Hercules was borrowed through Etruscan, where it is represented variously as Heracle, Hercle, and other forms. Hercules was a favorite subject for Etruscan art, and appears often on bronze mirrors. The Etruscan form Herceler derives from the Greek Heracles via syncope. A mild oath invoking Hercules (Hercule! or Mehercle!) was a common interjection in Classical Latin.

Hercules had a number of myths that were distinctly Roman. One of these is Hercules' defeat of Cacus, who was terrorizing the countryside of Rome. The hero was associated with the Aventine Hill through his son Aventinus. Mark Antony considered him a personal patron god, as did the emperor Commodus. Hercules received various forms of religious veneration, including as a deity concerned with children and childbirth, in part because of myths about his precocious infancy, and in part because he fathered countless children. Roman brides wore a special belt tied with the "knot of Hercules", which was supposed to be hard to untie. The comic playwright Plautus presents the myth of Hercules' conception as a sex comedy in his play Amphitryon; Seneca wrote the tragedy Hercules Furens about his bout with madness. During the Roman Imperial era, Hercules was worshipped locally from Hispania through Gaul.

Germanic association

Tacitus records a special affinity of the Germanic peoples for Hercules. In chapter 3 of his Germania, Tacitus states:

Some have taken this as Tacitus equating the Germanic Þunraz with Hercules by way of interpretatio romana.

In the Roman era Hercules' Club amulets appear from the 2nd to 3rd century, distributed over the empire (including Roman Britain, cf. Cool 1986), mostly made of gold, shaped like wooden clubs. A specimen found in Köln-Nippes bears the inscription "DEO HER [culi]", confirming the association with Hercules.

In the 5th to 7th centuries, during the Migration Period, the amulet is theorized to have rapidly spread from the Elbe Germanic area across Europe. These Germanic "Donar's Clubs" were made from deer antler, bone or wood, more rarely also from bronze or precious metals. The amulet type is replaced by the Viking Age Thor's hammer pendants in the course of the Christianization of Scandinavia from the 8th to 9th century.

Medieval mythography

After the Roman Empire became Christianized, mythological narratives were often reinterpreted as allegory, influenced by the philosophy of late antiquity. In the 4th century, Servius had described Hercules' return from the underworld as representing his ability to overcome earthly desires and vices, or the earth itself as a consumer of bodies. In medieval mythography, Hercules was one of the heroes seen as a strong role model who demonstrated both valor and wisdom, while the monsters he battles were regarded as moral obstacles. One glossator noted that when Hercules became a constellation, he showed that strength was necessary to gain entrance to Heaven.

Medieval mythography was written almost entirely in Latin, and original Greek texts were little used as sources for Hercules' myths.

Renaissance mythography

The Renaissance and the invention of the printing press brought a renewed interest in and publication of Greek literature. Renaissance mythography drew more extensively on the Greek tradition of Heracles, typically under the Romanized name Hercules, or the alternate name Alcides. In a chapter of his book Mythologiae (1567), the influential mythographer Natale Conti collected and summarized an extensive range of myths concerning the birth, adventures, and death of the hero under his Roman name Hercules. Conti begins his lengthy chapter on Hercules with an overview description that continues the moralizing impulse of the Middle Ages:

Hercules, who subdued and destroyed monsters, bandits, and criminals, was justly famous and renowned for his great courage. His great and glorious reputation was worldwide, and so firmly entrenched that he'll always be remembered. In fact the ancients honored him with his own temples, altars, ceremonies, and priests. But it was his wisdom and great soul that earned those honors; noble blood, physical strength, and political power just aren't good enough.

In 1600, the citizens of Avignon bestowed on Henry of Navarre (the future King Henry IV of France) the title of the Hercule Gaulois ("Gallic Hercules"), justifying the extravagant flattery with a genealogy that traced the origin of the House of Navarre to a nephew of Hercules' son Hispalus.

Road of Hercules
The Road of Hercules is a route across Southern Gaul that is associated with the path Hercules took during his 10th labor of retrieving the Cattle of Geryon from the Red Isles. Hannibal took the same path on his march towards Italy and encouraged the belief that he was the second Hercules. Primary sources often make comparisons between Hercules and Hannibal. Hannibal further tried to invoke parallels between himself and Hercules by starting his march on Italy by visiting the shrine of Hercules at Gades. While crossing the alps, he performed labors in a heroic manner. A famous example was noted by Livy, when Hannibal fractured the side of a cliff that was blocking his march.

Worship from women
In ancient Roman society women were usually limited to two types of cults: those that addressed feminine matters such as childbirth, and cults that required virginal chastity. However, there is evidence suggesting there were female worshippers of Apollo, Mars, Jupiter, and Hercules. Some scholars believe that women were completely prohibited from any of Hercules's cults. Others believe it was only the "Ara Maxima" at which they were not allowed to worship. Macrobius in his first book of Saturnalia paraphrases from Varro: "For when Hercules was bringing the cattle of Geryon through Italy, a woman replied to the thirsty hero that she could not give him water because it was the day of the Goddess Women and it was unlawful for a man to taste what had been prepared for her. Hercules, therefore, when he was about to offer a sacrifice forbid the presence of women and ordered Potitius and Pinarius who were in charge of his rites, not to allow any women from taking part". Macrobius states that women were restricted in their participation in Hercules cults, but to what extent remains ambiguous. He mentions that women were not allowed to participate in Sacrum which is general term used to describe anything that was believed to have belonged to the gods. This could include anything from a precious item to a temple. Due to the general nature of a Sacrum, we can not judge the extent of the prohibition from Macrobius alone. There are also ancient writings on this topic from Aulus Gellius when speaking on how Romans swore oaths. He mentioned that Roman women do not swear on Hercules, nor do Roman men swear on Castor. He went on to say that women refrain from sacrificing to Hercules. Propertius in his poem 4.9 also mentions similar information as Macrobius. This is evidence that he was also using Varro as a source.

Worship in myth
There is evidence of Hercules worship in myth in the Latin epic poem, the Aeneid. In the 8th book of the poem Aeneas finally reaches the future site of Rome, where he meets Evander and the Arcadians making sacrifices to Hercules on the banks of the Tiber river. They share a feast, and Evander tells the story of how Hercules defeated the monster Cascus, and describes him as a triumphant hero. Translated from the Latin text of Vergil, Evander stated: "Time brought to us in our time of need the aid and arrival of a god. For there came that mightiest avenger, the victor Hercules, proud with the slaughter and the spoils of threefold Geryon, and he drove the mighty bulls here, and the cattle filled both valley and riverside.

Hercules was also mentioned in the Fables of Gaius Julius Hyginus. For example, in his fable about Philoctetes he tells the story of how Philoctetes built a funeral pyre for Hercules so his body could be consumed and raised to immortality.

Hercules and the Roman triumph
According to Livy (9.44.16) Romans were commemorating military victories by building statues to Hercules as early as 305 BCE. Also, philosopher Pliny the Elder dates Hercules worship back to the time of Evander, by accrediting him with erecting a statue in the Forum Boarium of Hercules. Scholars agree that there would have been 5–7 temples in Augustan Rome. There are believed to be related Republican triumphatores, however, not necessarily triumphal dedications. There are two temples located in the Campus Martius. One, being the Temple of Hercules Musarum, dedicated between 187 and 179 BCE by M. Fulvius Nobilior. And the other being the Temple of Hercules Custos, likely renovated by Sulla in the 80s BCE.

In art
In Roman works of art and in Renaissance and post-Renaissance art, Hercules can be identified by his attributes, the lion skin and the gnarled club (his favorite weapon); in mosaic he is shown tanned bronze, a virile aspect.

In the twentieth century, the Farnese Hercules has inspired artists such as Jeff Koons, Matthew Darbyshire and Robert Mapplethorpe to reinterpret Hercules for new audiences. The choice of deliberately white materials by Koons and Darbyshire has been interpreted as perpetuation of colourism in how the classical world is viewed. Mapplethorpe's work with black model Derrick Cross can be seen as a reaction to Neo-classical colourism, resisting the portrayal of Hercules as white.

In numismatics
Hercules was among the earliest figures on ancient Roman coinage, and has been the main motif of many collector coins and medals since. One example is the Austrian 20 euro Baroque Silver coin issued on September 11, 2002. The obverse side of the coin shows the Grand Staircase in the town palace of Prince Eugene of Savoy in Vienna, currently the Austrian Ministry of Finance. Gods and demi-gods hold its flights, while Hercules stands at the turn of the stairs.

Military
Six successive ships of the British Royal Navy, from the 18th to the 20th century, bore the name HMS Hercules.

In the French Navy, there were no less than nineteen ships called Hercule, plus three more named Alcide which is another name of the same hero.

Hercules' name was also used for five ships of the US Navy, four ships of the Spanish Navy, four of the Argentine Navy and two of the Swedish Navy, as well as for numerous civilian sailing and steam ships.

In modern aviation a military transport aircraft produced by Lockheed Martin carries the title Lockheed C-130 Hercules.

Operation Herkules was the German code-name given to an abortive plan for the invasion of Malta during the Second World War.

In films
A series of nineteen Italian Hercules movies were made in the late 1950s and early 1960s. The actors who played Hercules in these films were Steve Reeves, Gordon Scott, Kirk Morris, Mickey Hargitay, Mark Forest, Alan Steel, Dan Vadis, Brad Harris, Reg Park, Peter Lupus (billed as Rock Stevens) and Michael Lane. A number of English-dubbed Italian films that featured the name of Hercules in their title were not intended to be movies about Hercules.

title: Hercules