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Fertilisation

Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or offspring. While processes such as insemination or pollination, which happen before the fusion of gametes, are also sometimes informally referred to as fertilisation, these are technically separate processes. The cycle of fertilisation and development of new individuals is called sexual reproduction. During double fertilisation in angiosperms, the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation.

History
In antiquity, Aristotle conceived the formation of new individuals through fusion of male and female fluids, with form and function emerging gradually, in a mode called by him as epigenetic.

In 1784, Spallanzani established the need of interaction between the female's ovum and male's sperm to form a zygote in frogs. In 1827, Karl Ernst von Baer observed a therian mammalian egg for the first time. Oscar Hertwig (1876), in Germany, described the fusion of nuclei of spermatozoa and of ova from sea urchin.

Evolution
The evolution of fertilisation is related to the origin of meiosis, as both are part of sexual reproduction, originated in eukaryotes. One hypothesis states that meiosis originated from mitosis.

Fertilisation in plants

The gametes that participate in fertilisation of plants are the sperm (male) and the egg (female) cell. Various plant groups have differing methods by which the gametes produced by the male and female gametophytes come together and are fertilised. In bryophytes and pteridophytic land plants, fertilisation of the sperm and egg takes place within the archegonium. In seed plants, the male gametophyte is formed within a pollen grain. After pollination, the pollen grain germinates, and a pollen tube grows and penetrates the ovule through a tiny pore called a micropyle. The sperm are transferred from the pollen through the pollen tube to the ovule where the egg is fertilised. In flowering plants, two sperm cells are released from the pollen tube, and a second fertilisation event occurs involving the second sperm cell and the central cell of the ovule, which is a second female gamete.

Pollen tube growth
Unlike animal sperm which is motile, the sperm of most seed plants is immotile and relies on the pollen tube to carry it to the ovule where the sperm is released. The pollen tube penetrates the stigma and elongates through the extracellular matrix of the style before reaching the ovary. Then near the receptacle, it breaks through the ovule through the micropyle (an opening in the ovule wall) and the pollen tube "bursts" into the embryo sac, releasing sperm. The growth of the pollen tube has been believed to depend on chemical cues from the pistil, however these mechanisms were poorly understood until 1995. Work done on tobacco plants revealed a family of glycoproteins called TTS proteins that enhanced growth of pollen tubes. Pollen tubes in a sugar free pollen germination medium and a medium with purified TTS proteins both grew. However, in the TTS medium, the tubes grew at a rate 3x that of the sugar-free medium. TTS proteins were also placed on various locations of semi in vivo pollinated pistils, and pollen tubes were observed to immediately extend toward the proteins. Transgenic plants lacking the ability to produce TTS proteins had slower pollen tube growth and reduced fertility.

Rupture of pollen tube
The rupture of the pollen tube to release sperm in Arabidopsis has been shown to depend on a signal from the female gametophyte. Specific proteins called FER protein kinases present in the ovule control the production of highly reactive derivatives of oxygen called reactive oxygen species (ROS). ROS levels have been shown via GFP to be at their highest during floral stages when the ovule is the most receptive to pollen tubes, and lowest during times of development and following fertilisation. High amounts of ROS activate Calcium ion channels in the pollen tube, causing these channels to take up Calcium ions in large amounts. This increased uptake of calcium causes the pollen tube to rupture, and release its sperm into the ovule. Pistil feeding assays in which plants were fed diphenyl iodonium chloride (DPI) suppressed ROS concentrations in Arabidopsis, which in turn prevented pollen tube rupture.

Flowering plants
After being fertilised, the ovary starts to swell and develop into the fruit. With multi-seeded fruits, multiple grains of pollen are necessary for syngamy with each ovule. The growth of the pollen tube is controlled by the vegetative (or tube) cytoplasm. Hydrolytic enzymes are secreted by the pollen tube that digest the female tissue as the tube grows down the stigma and style; the digested tissue is used as a nutrient source for the pollen tube as it grows. During pollen tube growth towards the ovary, the generative nucleus divides to produce two separate sperm nuclei (haploid number of chromosomes) – a growing pollen tube therefore contains three separate nuclei, two sperm and one tube. The sperms are interconnected and dimorphic, the large one, in a number of plants, is also linked to the tube nucleus and the interconnected sperm and the tube nucleus form the "male germ unit".

Double fertilisation is the process in angiosperms (flowering plants) in which two sperm from each pollen tube fertilise two cells in a female gametophyte (sometimes called an embryo sac) that is inside an ovule. After the pollen tube enters the gametophyte, the pollen tube nucleus disintegrates and the two sperm cells are released; one of the two sperm cells fertilises the egg cell (at the bottom of the gametophyte near the micropyle), forming a diploid (2n) zygote. This is the point when fertilisation actually occurs; pollination and fertilisation are two separate processes. The nucleus of the other sperm cell fuses with two haploid polar nuclei (contained in the central cell) in the centre of the gametophyte. The resulting cell is triploid (3n). This triploid cell divides through mitosis and forms the endosperm, a nutrient-rich tissue, inside the seed. The two central-cell maternal nuclei (polar nuclei) that contribute to the endosperm arise by mitosis from the single meiotic product that also gave rise to the egg. Therefore, maternal contribution to the genetic constitution of the triploid endosperm is double that of the embryo.

One primitive species of flowering plant, Nuphar polysepala, has endosperm that is diploid, resulting from the fusion of a sperm with one, rather than two, maternal nuclei. It is believed that early in the development of angiosperm lineages, there was a duplication in this mode of reproduction, producing seven-celled/eight-nucleate female gametophytes, and triploid endosperms with a 2:1 maternal to paternal genome ratio.

In many plants, the development of the flesh of the fruit is proportional to the percentage of fertilised ovules. For example, with watermelon, about a thousand grains of pollen must be delivered and spread evenly on the three lobes of the stigma to make a normal sized and shaped fruit.

Self-pollination and outcrossing
Outcrossing, or cross-fertilisation, and self-fertilisation represent different strategies with differing benefits and costs. An estimated 48.7% of plant species are either dioecious or self-incompatible obligate outcrossers. It is also estimated that about 42% of flowering plants exhibit a mixed mating system in nature.

In the most common kind of mixed mating system, individual plants produce a single type of flower and fruits may contain self-fertilised, outcrossed or a mixture of progeny types. The transition from cross-fertilisation to self-fertilisation is the most common evolutionary transition in plants, and has occurred repeatedly in many independent lineages. About 10-15% of flowering plants are predominantly self-fertilising.

Under circumstances where pollinators or mates are rare, self-fertilisation offers the advantage of reproductive assurance. Self-fertilisation can therefore result in improved colonisation ability. In some species, self-fertilisation has persisted over many generations. Capsella rubella is a self-fertilising species that became self-compatible 50,000 to 100,000 years ago. Arabidopsis thaliana is a predominantly self-fertilising plant with an out-crossing rate in the wild of less than 0.3%; a study suggested that self-fertilisation evolved roughly a million years ago or more in A. thaliana. In long-established self-fertilising plants, the masking of deleterious mutations and the production of genetic variability is infrequent and thus unlikely to provide a sufficient benefit over many generations to maintain the meiotic apparatus. Consequently, one might expect self-fertilisation to be replaced in nature by an ameiotic asexual form of reproduction that would be less costly. However the actual persistence of meiosis and self-fertilisation as a form of reproduction in long-established self-fertilising plants may be related to the immediate benefit of efficient recombinational repair of DNA damage during formation of germ cells provided by meiosis at each generation.

Fertilisation in animals
The mechanics behind fertilisation has been studied extensively in sea urchins and mice. This research addresses the question of how the sperm and the appropriate egg find each other and the question of how only one sperm gets into the egg and delivers its contents. There are three steps to fertilisation that ensure species-specificity:

 * 1) Chemotaxis
 * 2) Sperm activation/acrosomal reaction
 * 3) Sperm/egg adhesion

Internal vs. external
Consideration as to whether an animal (more specifically a vertebrate) uses internal or external fertilisation is often dependent on the method of birth. Oviparous animals laying eggs with thick calcium shells, such as chickens, or thick leathery shells generally reproduce via internal fertilisation so that the sperm fertilises the egg without having to pass through the thick, protective, tertiary layer of the egg. Ovoviviparous and viviparous animals also use internal fertilisation. Although some organisms reproduce via amplexus, they may still use internal fertilisation, as with some salamanders. Advantages of internal fertilisation include minimal waste of gametes, greater chance of individual egg fertilisation, longer period of egg protection, and selective fertilisation. Many females have the ability to store sperm for extended periods of time and can fertilise their eggs at their own desire.

Oviparous animals producing eggs with thin tertiary membranes or no membranes at all, on the other hand, use external fertilisation methods. Such animals may be more precisely termed ovuliparous. External fertilisation is advantageous in that it minimises contact (which decreases the risk of disease transmission), and greater genetic variation.

Sea urchins

Sperm find the eggs via chemotaxis, a type of ligand/receptor interaction. Resact is a 14 amino acid peptide purified from the jelly coat of A. punctulata that attracts the migration of sperm.

After finding the egg, the sperm penetrates the jelly coat through a process called sperm activation. In another ligand/receptor interaction, an oligosaccharide component of the egg binds and activates a receptor on the sperm and causes the acrosomal reaction. The acrosomal vesicles of the sperm fuse with the plasma membrane and are released. In this process, molecules bound to the acrosomal vesicle membrane, such as bindin, are exposed on the surface of the sperm. These contents digest the jelly coat and eventually the vitelline membrane. In addition to the release of acrosomal vesicles, there is explosive polymerisation of actin to form a thin spike at the head of the sperm called the acrosomal process.

The sperm binds to the egg through another ligand reaction between receptors on the vitelline membrane. The sperm surface protein bindin, binds to a receptor on the vitelline membrane identified as EBR1.

Fusion of the plasma membranes of the sperm and egg are likely mediated by bindin. At the site of contact, fusion causes the formation of a fertilisation cone.

Mammals
Mammals internally fertilise through copulation. After a male ejaculates, many sperm move to the upper vagina (via contractions from the vagina) through the cervix and across the length of the uterus to meet the ovum. In cases where fertilisation occurs, the female usually ovulates during a period that extends from hours before copulation to a few days after; therefore, in most mammals, it is more common for ejaculation to precede ovulation than vice versa.

When sperm are deposited into the anterior vagina, they are not capable of fertilisation (i.e., non-capacitated) and are characterised by slow linear motility patterns. This motility, combined with muscular contractions enables sperm transport towards the uterus and oviducts. There is a pH gradient within the micro-environment of the female reproductive tract such that the pH near the vaginal opening is lower (approximately 5) than the oviducts (approximately 8). The sperm-specific pH-sensitive calcium transport protein called CatSper increases the sperm cell permeability to calcium as it moves further into the reproductive tract. Intracellular calcium influx contributes to sperm capacitation and hyperactivation, causing a more violent and rapid non-linear motility pattern as sperm approach the oocyte. The capacitated spermatozoon and the oocyte meet and interact in the ampulla of the fallopian tube. Rheotaxis, thermotaxis and chemotaxis are known mechanisms that guide sperm towards the egg during the final stage of sperm migration. Spermatozoa respond (see Sperm thermotaxis) to the temperature gradient of ~2 °C between the oviduct and the ampulla, and chemotactic gradients of progesterone have been confirmed as the signal emanating from the cumulus oophorus cells surrounding rabbit and human oocytes. Capacitated and hyperactivated sperm respond to these gradients by changing their behaviour and moving towards the cumulus-oocyte complex. Other chemotactic signals such as formyl Met-Leu-Phe (fMLF) may also guide spermatozoa.

The zona pellucida, a thick layer of extracellular matrix that surrounds the egg and is similar to the role of the vitelline membrane in sea urchins, binds the sperm. Unlike sea urchins, the sperm binds to the egg before the acrosomal reaction. ZP3, a glycoprotein in the zona pellucida, is responsible for egg/sperm adhesion in humans. The receptor galactosyltransferase (GalT) binds to the N-acetylglucosamine residues on the ZP3 and is important for binding with the sperm and activating the acrosome reaction. ZP3 is sufficient though unnecessary for sperm/egg binding. Two additional sperm receptors exist: a 250kD protein that binds to an oviduct secreted protein, and SED1, which independently binds to the zona. After the acrosome reaction, the sperm is believed to remain bound to the zona pellucida through exposed ZP2 receptors. These receptors are unknown in mice but have been identified in guinea pigs.

In mammals, the binding of the spermatozoon to the GalT initiates the acrosome reaction. This process releases the hyaluronidase that digests the matrix of hyaluronic acid in the vestments around the oocyte. Additionally, heparin-like glycosaminoglycans (GAGs) are released near the oocyte that promote the acrosome reaction. Fusion between the oocyte plasma membranes and sperm follows and allows the sperm nucleus, the typical centriole, and atypical centriole that is attached to the flagellum, but not the mitochondria, to enter the oocyte. The protein CD9 likely mediates this fusion in mice (the binding homolog). The egg "activates" itself upon fusing with a single sperm cell and thereby changes its cell membrane to prevent fusion with other sperm. Zinc atoms are released during this activation.

This process ultimately leads to the formation of a diploid cell called a zygote. The zygote divides to form a blastocyst and, upon entering the uterus, implants in the endometrium, beginning pregnancy. Embryonic implantation not in the uterine wall results in an ectopic pregnancy that can kill the mother.

In such animals as rabbits, coitus induces ovulation by stimulating the release of the pituitary hormone gonadotropin; this release greatly increases the likelihood of pregnancy.

Humans

Fertilisation in humans is the union of a human egg and sperm, usually occurring in the ampulla of the fallopian tube, producing a single celled zygote, the first stage of life in the development of a genetically unique organism, and initiating embryonic development. Scientists discovered the dynamics of human fertilisation in the nineteenth century.

The term conception commonly refers to "the process of becoming pregnant involving fertilisation or implantation or both". Its use makes it a subject of semantic arguments about the beginning of pregnancy, typically in the context of the abortion debate.

Upon gastrulation, which occurs around 16 days after fertilisation, the implanted blastocyst develops three germ layers, the endoderm, the ectoderm and the mesoderm, and the genetic code of the father becomes fully involved in the development of the embryo; later twinning is impossible. Additionally, interspecies hybrids survive only until gastrulation and cannot further develop. However, some human developmental biology literature refers to the conceptus and such medical literature refers to the "products of conception" as the post-implantation embryo and its surrounding membranes. The term "conception" is not usually used in scientific literature because of its variable definition and connotation.

Insects

Insects in different groups, including the Odonata (dragonflies and damselflies) and the Hymenoptera (ants, bees, and wasps) practise delayed fertilisation. Among the Odonata, females may mate with multiple males, and store sperm until the eggs are laid. The male may hover above the female during egg-laying (oviposition) to prevent her from mating with other males and replacing his sperm; in some groups such as the darters, the male continues to grasp the female with his claspers during egg-laying, the pair flying around in tandem. Among social Hymenoptera, honeybee queens mate only on mating flights, in a short period lasting some days; a queen may mate with eight or more drones. She then stores the sperm for the rest of her life, perhaps for five years or more.

Fertilisation in fungi
In many fungi (except chytrids), as in some protists, fertilisation is a two step process. First, the cytoplasms of the two gamete cells fuse (called plasmogamy), producing a dikaryotic or heterokaryotic cell with multiple nuclei. This cell may then divide to produce dikaryotic or heterokaryotic hyphae. The second step of fertilisation is karyogamy, the fusion of the nuclei to form a diploid zygote.

In chytrid fungi, fertilisation occurs in a single step with the fusion of gametes, as in animals and plants.

Fertilisation in protozoa
There are three types of fertilisation processes in protozoa:
 * gametogamy;
 * autogamy;
 * gamontogamy.

Fertilisation in algae
Algae, like some land plants, undergo alternation of generations. Some algae are isomorphic, where both the sporophyte (2n) and gameteophyte (n) are the same morphologically. When algae reproduction is described as oogamous, the male and female gametes are different morphologically, where there is a large non-motile egg for female gametes, and the male gamete are uniflagellate (motile). Via the process of syngamy, these will form a new zygote, regenerating the sporophyte generation again.

Fertilisation and genetic recombination
Meiosis results in a random segregation of the genes that each parent contributes. Each parent organism is usually identical save for a fraction of their genes; each gamete is therefore genetically unique. At fertilisation, parental chromosomes combine. In humans, (2²²)² = 17.6x1012 chromosomally different zygotes are possible for the non-sex chromosomes, even assuming no chromosomal crossover. If crossover occurs once, then on average (4²²)² = 309x1024 genetically different zygotes are possible for every couple, not considering that crossover events can take place at most points along each chromosome. The X and Y chromosomes undergo no crossover events and are therefore excluded from the calculation. The mitochondrial DNA is only inherited from the maternal parent.

The sperm aster and zygote centrosomes
Shortly after the sperm fuse with the egg, the two sperm centrioles form the embryo first centrosome and microtubule aster. The sperm centriole, found near the male pronucleus, recruit egg Pericentriolar material proteins forming the zygote first centrosome. This centrosome nucleates microtubules in the shape of stars called astral microtubules. The microtubules span the whole valium of the egg, allowing the egg pronucleus to use the cables to get to the male pronucleus. As the male and female pronuclei approach each other, the single centrosome split into two centrosomes located in the interphase between the pronuclei. Then the centrosome via the astral microtubules polarises the genome inside the pronuclei.

Parthenogenesis
Organisms that normally reproduce sexually can also reproduce via parthenogenesis, wherein an unfertilised female gamete produces viable offspring. These offspring may be clones of the mother, or in some cases genetically differ from her but inherit only part of her DNA. Parthenogenesis occurs in many plants and animals and may be induced in others through a chemical or electrical stimulus to the egg cell. In 2004, Japanese researchers led by Tomohiro Kono succeeded after 457 attempts to merge the ova of two mice by blocking certain proteins that would normally prevent the possibility; the resulting embryo normally developed into a mouse.

Allogamy and autogamy
Allogamy, which is also known as cross-fertilisation, refers to the fertilisation of an egg cell from one individual with the male gamete of another.

Autogamy which is also known as self-fertilisation, occurs in such hermaphroditic organisms as plants and flatworms; therein, two gametes from one individual fuse.

Other variants of bisexual reproduction
Some relatively unusual forms of reproduction are:

Gynogenesis: A sperm stimulates the egg to develop without fertilisation or syngamy. The sperm may enter the egg.

Hybridogenesis: One genome is eliminated to produce haploid eggs.

Canina meiosis: (sometimes called "permanent odd polyploidy") one genome is transmitted in the Mendelian fashion, others are transmitted clonally.

Benefits of cross-fertilisation
The major benefit of cross-fertilisation is generally thought to be the avoidance of inbreeding depression. Charles Darwin, in his 1876 book The Effects of Cross and Self Fertilisation in the Vegetable Kingdom (pages 466-467) summed up his findings in the following way. "'It has been shown in the present volume that the offspring from the union of two distinct individuals, especially if their progenitors have been subjected to very different conditions, have an immense advantage in height, weight, constitutional vigour and fertility over the self-fertilised offspring from one of the same parents. And this fact is amply sufficient to account for the development of the sexual elements, that is, for the genesis of the two sexes.'"

In addition, it is thought by some, that a long-term advantage of out-crossing in nature is increased genetic variability that promotes adaptation or avoidance of extinction (see Genetic variability).

title: Fertilisation


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The Federalist Papers

The Federalist Papers is a collection of 85 articles and essays written by Alexander Hamilton, James Madison, and John Jay under the collective pseudonym "Publius" to promote the ratification of the Constitution of the United States. The collection was commonly known as The Federalist until the name The Federalist Papers emerged in the twentieth century.

The first seventy-seven of these essays were published serially in the Independent Journal, the New York Packet, and The Daily Advertiser between October 1787 and April 1788. A compilation of these 77 essays and eight others were published in two volumes as The Federalist: A Collection of Essays, Written in Favour of the New Constitution, as Agreed upon by the Federal Convention, September 17, 1787, by publishing firm J. & A. McLean in March and May 1788. The last eight papers (Nos. 78–85) were republished in the New York newspapers between June 14 and August 16, 1788.

The authors of The Federalist intended to influence the voters to ratify the Constitution. In Federalist No. 1, they explicitly set that debate in broad political terms:"It has been frequently remarked, that it seems to have been reserved to the people of this country, by their conduct and example, to decide the important question, whether societies of men are really capable or not, of establishing good government from reflection and choice, or whether they are forever destined to depend, for their political constitutions, on accident and force."

In Federalist No. 10, Madison discusses the means of preventing rule by majority faction and advocates a large, commercial republic. This is complemented by Federalist No. 14, in which Madison takes the measure of the United States, declares it appropriate for an extended republic, and concludes with a memorable defense of the constitutional and political creativity of the Federal Convention.

In Federalist No. 84, Hamilton makes the case that there is no need to amend the Constitution by adding a Bill of Rights, insisting that the various provisions in the proposed Constitution protecting liberty amount to a "bill of rights." Federalist No. 78, also written by Hamilton, lays the groundwork for the doctrine of judicial review by federal courts of federal legislation or executive acts. Federalist No. 70 presents Hamilton's case for a one-man chief executive. In Federalist No. 39, Madison presents the clearest exposition of what has come to be called "Federalism". In Federalist No. 51, Madison distills arguments for checks and balances in an essay often quoted for its justification of government as "the greatest of all reflections on human nature." According to historian Richard B. Morris, the essays that make up The Federalist Papers are an "incomparable exposition of the Constitution, a classic in political science unsurpassed in both breadth and depth by the product of any later American writer."

On June 21, 1788, the proposed Constitution was ratified by the minimum of nine states required under Article VII. In late July 1788, with eleven states having ratified the new Constitution, the process of organizing the new government began.

Origins

The Federal Convention (Constitutional Convention) sent the proposed Constitution to the Confederation Congress, which in turn submitted it to the states for ratification at the end of September 1787. On September 27, 1787, "Cato" first appeared in the New York press criticizing the proposition; "Brutus" followed on October 18, 1787. These and other articles and public letters critical of the new Constitution would eventually become known as the "Anti-Federalist Papers". In response, Alexander Hamilton decided to launch a measured defense and extensive explanation of the proposed Constitution to the people of the state of New York. He wrote in Federalist No. 1 that the series would "endeavor to give a satisfactory answer to all the objections which shall have made their appearance, that may seem to have any claim to your attention."

Hamilton recruited collaborators for the project. He enlisted John Jay, who after four essays (Federalist Nos. 2, 3, 4, and 5), fell ill and contributed only one more essay, Federalist No. 64, to the series. Jay also distilled his case into a pamphlet in the spring of 1788, An Address to the People of the State of New-York; Hamilton cited it approvingly in Federalist No. 85. James Madison, present in New York as a Virginia delegate to the Confederation Congress, was recruited by Hamilton and Jay and became Hamilton's primary collaborator. Gouverneur Morris and William Duer were also considered. However, Morris turned down the invitation, and Hamilton rejected three essays written by Duer. Duer later wrote in support of the three Federalist authors under the name "Philo-Publius", meaning either "Friend of the People" or "Friend of Hamilton" based on Hamilton's pen name Publius.

Hamilton chose the pseudonymous name "Publius". While many other pieces representing both sides of the constitutional debate were written under Roman names, historian Albert Furtwangler contends that Publius' was a cut above 'Caesar' or 'Brutus' or even 'Cato'. Publius Valerius helped found the ancient republic of Rome. His more famous name, Publicola, meant 'friend of the people'." Hamilton had applied this pseudonym to three letters in 1778, in which he attacked fellow Federalist Samuel Chase and revealed that Chase had taken advantage of knowledge gained in Congress to try to dominate the flour market.

Authorship

At the time of publication, the authors of The Federalist Papers attempted to hide their identities due to Hamilton and Madison having attended the convention. Astute observers, however, correctly discerned the identities of Hamilton, Madison, and Jay. Establishing authorial authenticity of the essays that constitute The Federalist Papers has not always been clear. After Hamilton's death in 1804, a list emerged, claiming that he alone had written two-thirds of The Federalist essays. Some believe that several of these essays were written by Madison (Nos. 49–58 and 62–63). The scholarly detective work of Douglass Adair in 1944 postulated the following assignments of authorship, corroborated in 1964 by a computer analysis of the text:

 * Alexander Hamilton (51 articles: Nos. 1, 6–9, 11–13, 15–17, 21–36, 59–61, and 65–85)
 * James Madison (29 articles: Nos. 10, 14, 18–20, 37–58 and 62–63)
 * John Jay (5 articles: Nos. 2–5 and 64).

In six months, a total of 85 articles were written by the three men. Hamilton, who had been a leading advocate of national constitutional reform throughout the 1780s and was one of the three representatives for New York at the Constitutional Convention, in 1789 became the first secretary of the treasury, a post he held until his resignation in 1795. Madison, who is now acknowledged as the father of the Constitution — despite his repeated rejection of this honor during his lifetime, became a leading member of the U.S. House of Representatives from Virginia (1789–1797), secretary of state (1801–1809), and ultimately the fourth president of the United States (1809–1817).

John Jay, who had been secretary for foreign affairs under the Articles of Confederation from 1784 through their expiration in 1789, became the first Chief Justice of the United States in 1789, stepping down in 1795 to accept election as governor of New York, a post he held for two terms, retiring in 1801.

Publication

The Federalist articles appeared in three New York newspapers: The Independent Journal, the New-York Packet, and the Daily Advertiser, beginning on October 27, 1787. Although written and published with haste, The Federalist articles were widely read and greatly influenced the shape of American political institutions. Hamilton, Madison and Jay published the essays at a rapid pace. At times, three to four new essays by Publius appeared in the papers in a single week. Garry Wills observes that this fast pace of production "overwhelmed" any possible response: "Who, given ample time could have answered such a battery of arguments? And no time was given." Hamilton also encouraged the reprinting of the essays in newspapers outside New York state, and indeed they were published in several other states where the ratification debate was taking place. However, they were only irregularly published outside New York, and in other parts of the country they were often overshadowed by local writers.

Because the essays were initially published in New York, most of them begin with the same salutation: "To the People of the State of New York".

The high demand for the essays led to their publication in a more permanent form. On January 1, 1788, the New York publishing firm J. & A. McLean announced that they would publish the first 36 essays as a bound volume; that volume was released on March 22, 1788, and was titled The Federalist Volume 1. New essays continued to appear in the newspapers; Federalist No. 77 was the last number to appear first in that form, on April 2. A second bound volume was released on May 28, containing Federalist Nos. 37–77 and the previously unpublished Nos. 78–85. The last eight papers (Nos. 78–85) were republished in the New York newspapers between June 14 and August 16, 1788.

A 1792 French edition ended the collective anonymity of Publius, announcing that the work had been written by "Mm. Hamilton, Maddisson e Gay, citoyens de l'État de New York". In 1802, George Hopkins published an American edition that similarly named the authors. Hopkins wished as well that "the name of the writer should be prefixed to each number," but at this point Hamilton insisted that this was not to be, and the division of the essays among the three authors remained a secret.

The first publication to divide the papers in such a way was an 1810 edition that used a list left by Hamilton to associate the authors with their numbers; this edition appeared as two volumes of the compiled "Works of Hamilton". In 1818, Jacob Gideon published a new edition with a new listing of authors, based on a list provided by Madison. The difference between Hamilton's list and Madison's formed the basis for a dispute over the authorship of a dozen of the essays.

Both Hopkins's and Gideon's editions incorporated significant edits to the text of the papers themselves, generally with the approval of the authors. In 1863, Henry Dawson published an edition containing the original text of the papers, arguing that they should be preserved as they were written in that particular historical moment, not as edited by the authors years later.

Modern scholars generally use the text prepared by Jacob E. Cooke for his 1961 edition of The Federalist; this edition used the newspaper texts for essay numbers 1–76 and the McLean edition for essay numbers 77–85.

Disputed essays

While the authorship of 73 of The Federalist essays is fairly certain, the identities of those who wrote the twelve remaining essays are disputed by some scholars. The modern consensus is that Madison wrote essays Nos. 49–58, with Nos. 18–20 being products of a collaboration between him and Hamilton; No. 64 was by John Jay. The first open designation of which essay belonged to whom was provided by Hamilton who, in the days before his ultimately fatal gun duel with Aaron Burr, provided his lawyer with a list detailing the author of each number. This list credited Hamilton with a full 63 of the essays (three of those being jointly written with Madison), almost three-quarters of the whole, and was used as the basis for an 1810 printing that was the first to make specific attribution for the essays.

Madison did not immediately dispute Hamilton's list, but provided his own list for the 1818 Gideon edition of The Federalist. Madison claimed 29 essays for himself, and he suggested that the difference between the two lists was "owing doubtless to the hurry in which [Hamilton's] memorandum was made out." A known error in Hamilton's list—Hamilton incorrectly ascribed No. 54 to John Jay, when in fact, Jay wrote No. 64—provided some evidence for Madison's suggestion.

Statistical analysis has been undertaken on several occasions in attempts to accurately identify the author of each individual essay. After examining word choice and writing style, studies generally agree that the disputed essays were written by James Madison. However, there are notable exceptions maintaining that some of the essays which are now widely attributed to Madison were, in fact, collaborative efforts.

Influence on the ratification debates
The Federalist Papers were written to support the ratification of the Constitution, specifically in New York. Whether they succeeded in this mission is questionable. Separate ratification proceedings took place in each state, and the essays were not reliably reprinted outside of New York; furthermore, by the time the series was well underway, a number of important states had already ratified it, for instance Pennsylvania on December 12. New York held out until July 26; certainly The Federalist was more important there than anywhere else, but Furtwangler argues that it "could hardly rival other major forces in the ratification contests"—specifically, these forces included the personal influence of well-known Federalists, for instance Hamilton and Jay, and Anti-Federalists, including Governor George Clinton. Further, by the time New York came to a vote, ten states had already ratified the Constitution and it had thus already passed—only nine states had to ratify it for the new government to be established among them; the ratification by Virginia, the tenth state, placed pressure on New York to ratify. In light of that, Furtwangler observes, "New York's refusal would make that state an odd outsider."

Only 19 Federalists were elected to New York's ratification convention, compared to the Anti-Federalists' 46 delegates. While New York did indeed ratify the Constitution on July 26, the lack of public support for pro-Constitution Federalists has led historian John Kaminski to suggest that the impact of The Federalist on New York citizens was "negligible".

As for Virginia, which ratified the Constitution only at its convention on June 25, Hamilton writes in a letter to Madison that the collected edition of The Federalist had been sent to Virginia; Furtwangler presumes that it was to act as a "debater's handbook for the convention there", though he claims that this indirect influence would be a "dubious distinction". Probably of greater importance to the Virginia debate, in any case, were George Washington's support for the proposed Constitution and the presence of Madison and Edmund Randolph, the governor, at the convention arguing for ratification.

Structure and content
In Federalist No. 1, Hamilton listed six topics to be covered in the subsequent articles:

 * 1) "The utility of the UNION to your political prosperity"—covered in No. 2 through No. 14
 * 2) "The insufficiency of the present Confederation to preserve that Union"—covered in No. 15 through No. 22
 * 3) "The necessity of a government at least equally energetic with the one proposed to the attainment of this object"—covered in No. 23 through No. 36
 * 4) "The conformity of the proposed constitution to the true principles of republican government"—covered in No. 37 through No. 84
 * 5) "Its analogy to your own state constitution"—covered in No. 85
 * 6) "The additional security which its adoption will afford to the preservation of that species of government, to liberty and to prosperity"—covered in No. 85.

Furtwangler notes that as the series grew, this plan was somewhat changed. The fourth topic expanded into detailed coverage of the individual articles of the Constitution and the institutions it mandated, while the two last topics were merely touched on in the last essay.

The papers can be broken down by author as well as by topic. At the start of the series, all three authors were contributing; the first 20 papers are broken down as 11 by Hamilton, five by Madison and four by Jay. The rest of the series, however, is dominated by three long segments by a single writer: Nos. 21–36 by Hamilton, Nos. 37–58 by Madison, written while Hamilton was in Albany, and No. 65 through the end by Hamilton, published after Madison had left for Virginia.

Opposition to the Bill of Rights
The Federalist Papers (specifically Federalist No. 84) are notable for their opposition to what later became the United States Bill of Rights. The idea of adding a Bill of Rights to the Constitution was originally controversial because the Constitution, as written, did not specifically enumerate or protect the rights of the people, rather it listed the powers of the government and left all that remained to the states and the people. Alexander Hamilton, the author of Federalist No. 84, feared that such an enumeration, once written down explicitly, would later be interpreted as a list of the only rights that people had.

However, Hamilton's opposition to a Bill of Rights was far from universal. Robert Yates, writing under the pseudonym "Brutus", articulated this view point in the so-called Anti-Federalist No. 84, asserting that a government unrestrained by such a bill could easily devolve into tyranny. References in The Federalist and in the ratification debates warn of demagogues of the variety who through divisive appeals would aim at tyranny. The Federalist begins and ends with this issue. In the final paper Hamilton offers "a lesson of moderation to all sincere lovers of the Union, and ought to put them on their guard against hazarding anarchy, civil war, a perpetual alienation of the States from each other, and perhaps the military despotism of a successful demagogue". The matter was further clarified by the Ninth Amendment.

Judicial use
Federal judges, when interpreting the Constitution, frequently use The Federalist Papers as a contemporary account of the intentions of the framers and ratifiers. They have been applied on issues ranging from the power of the federal government in foreign affairs (in Hines v. Davidowitz) to the validity of ex post facto laws (in the 1798 decision Calder v. Bull, apparently the first decision to mention The Federalist). , The Federalist had been quoted 291 times in Supreme Court decisions.

The amount of deference that should be given to The Federalist Papers in constitutional interpretation has always been somewhat controversial. As early as 1819, Chief Justice John Marshall noted in the famous case McCulloch v. Maryland, that "the opinions expressed by the authors of that work have been justly supposed to be entitled to great respect in expounding the Constitution. No tribute can be paid to them which exceeds their merit; but in applying their opinions to the cases which may arise in the progress of our government, a right to judge of their correctness must be retained." In a letter to Thomas Ritchie in 1821, James Madison stated of the Constitution that "the legitimate meaning of the Instrument must be derived from the text itself; or if a key is to be sought elsewhere, it must be not in the opinions or intentions of the Body which planned & proposed the Constitution, but in the sense attached to it by the people in their respective State Conventions where it recd. all the authority which it possesses."

Complete list
The colors used to highlight the rows correspond to the author of the paper.

{{legend|#BFE4FF|Alexander Hamilton}} {{legend|#FFDFBF|John Jay}} {{legend|#FFBFBF|James Madison}}

In popular culture
The purposes and authorship of The Federalist Papers were prominently highlighted in the lyrics of "Non-Stop", the finale of Act One in the 2015 Broadway musical Hamilton, written by Lin-Manuel Miranda.

General and cited references

 * Updated 2nd ed., originally published as
 * Wills, Gary. Explaining America: The Federalist. Garden City, NJ: 1981. ISBN 978-0140298390.
 * Wills, Gary. Explaining America: The Federalist. Garden City, NJ: 1981. ISBN 978-0140298390.
 * Wills, Gary. Explaining America: The Federalist. Garden City, NJ: 1981. ISBN 978-0140298390.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: The Federalist Papers


text:
Cisco (disambiguation)

Cisco is an American technology conglomerate.

Cisco may also refer to:

Places

 * Siska, British Columbia, also known historically as Cisco

United States

 * Cisco, California, a small town
 * Cisco, Georgia, an unincorporated community
 * Cisco, Illinois, a village
 * Cisco, Kentucky, an unincorporated community
 * Cisco, Minnesota, a former unincorporated community
 * Lisco, Nebraska, a small town in western Nebraska, erroneously called "Cisco" by the United States Census Bureau in 2000
 * Cisco, Texas, a city
 * Cisco Group, a geologic group in Texas
 * Cisco, Utah, a ghost town
 * Cisco Oil Field, Utah
 * Cisco, West Virginia, an unincorporated community

Companies

 * Certis CISCO, corporatised entity of the former Commercial and Industrial Security Corporation in Singapore
 * Cisco Brewers, a brewery, distillery and winery on Nantucket Island in Massachusetts, US
 * Nissin Cisco Co. Ltd, a Japanese cereal and snack-food manufacturer, subsidiary of Nissin Foods
 * Luzon Cisco Transport, Inc., a bus company which become part of Five Star Bus Company

Other uses

 * Cisco (name), a given name, nickname or surname (and a list of people with the name)
 * Cisco (fish), certain Eurasian and North American salmonid freshwater whitefish in the genus Coregonus
 * Cisco (Grand Theft Auto Advance), a character from Grand Theft Auto Advance
 * USS Cisco (SS-290), World War II submarine
 * Don Cisco or Cisco, Mexican-American rapper and producer
 * Cisco College, a community college in Cisco, Texas, US
 * Cisco Field, a proposed baseball stadium for the Oakland Athletics in California, US
 * Cisco High School (Texas)
 * Cisco (wine), a brand of low-end fortified wine
 * Cisco, an original composition performed on Pat Martino's 1967 album El_Hombre

title: Cisco (disambiguation)


text:
Federalist

The term federalist describes several political beliefs around the world. It may also refer to the concept of parties, whose members or supporters call themselves Federalists.

Europe federation
In Europe, proponents of deeper European integration are sometimes called Federalists. A major European NGO and advocacy group campaigning for such a political union is the Union of European Federalists. Movements towards a peacefully unified European state have existed since the 1920s, notably the Paneuropean Union. A pan-European party with representation in the European Parliament fighting for the same cause is Volt Europa.

In the European Parliament the Spinelli Group brings together MEPs from different political groups to work together of ideas and projects of European federalism; taking their name from Italian politician and MEP Altiero Spinelli, who himself was a major proponent of European federalism, also meeting with fellow deputies in the Crocodile Club.

Notable European Federalists are former European Commission president Jean-Claude Juncker, current EC president Ursula von der Leyen, leader of ALDE group Guy Verhofstadt, German Federal Minister for Economic Affairs and Energy of Germany Peter Altmaier, German MEP Elmar Brok and the former leader of the SPD Martin Schulz.

Latin America
In the Spanish-speaking parts of Latin America the term "federalist" is used in reference to the politics of 19th-century Argentina and Colombia. The Federalists opposed the Unitarians in Argentina and the Centralists in Colombia through the 19th century. Federalists fought for complete self-government and full provincial autonomy, as opposed to the centralized government that the Unitarians and Centralists favored. Furthermore, Federalists demanded tariff protection for their industries and, in Argentina, called for the end of the Buenos Aires customs as the only intermediary for foreign trade. During the Federal War (1859-1863) in Venezuela, liberal caudillos confronted conservatives, leading to the establishment of the modern federal States of Venezuela.

Argentina
The one Federalist leader in the Platine Region was José Gervasio Artigas, who opposed the centralist governments in Buenos Aires that followed the May Revolution, and created instead the Federal League in 1814 among several Argentine Provinces and the Banda Oriental (modern-day Uruguay). In 1819, the Federal armies rejected the centralist Constitution of the United Provinces of South America and defeated the forces of Supreme Director José Rondeau at the 1820 Battle of Cepeda, effectively ending the central government and securing Provinces' sovereignty through a series of inter-Provincial pacts (v.g. Treaty of Pilar, Treaty of Benegas, Quadrilateral Treaty). A new National Constitution was proposed only in 1826, during the Presidency of Unitarian Bernardino Rivadavia, but it was again rejected by the Provinces, leading to the dissolution of the National Government the following year.

Federalist Buenos Aires Governor Manuel Dorrego took over the management of the foreign affairs of the United Provinces, but he was deposed and executed in 1828 by Unitarian General Juan Lavalle, who commanded troops dissatisfied with the negotiations that ended the War with Brazil. The following year, Juan Manuel de Rosas, leader of Buenos Aires Federalists, defeated Lavalle and secured his resignation. Rosas was elected Governor of Buenos Aires later that year by the Provincial Legislature. To counteract these developments, the Unitarian League was created by General José María Paz in 1830, uniting nine Argentine Provinces. The 1831 Federal Pact between Buenos Aires, Entre Ríos and Santa Fe Provinces opposed a military alliance to the League and ultimately defeated it during 1832, its former members joining the Federal Pact into a loose confederation of Provinces known as the Argentine Confederation. Although the Unitarians were exiled in neighboring countries, the Civil War continued for two decades.

Buenos Aires Governor Juan Manuel de Rosas exerted a growing hegemony over the rest of the country during his 1835-1852 Government and resisted several Unitarian uprisings, but was finally defeated in 1852 by a coalition Army gathered by Entre Ríos Federalist Governor Justo José de Urquiza, who accused Rosas of not complying with Federal Pact provisions for a National Constitution. In 1853, a Federal Constitution was enacted (the current Constitution of Argentina, through amendments) and Urquiza was elected President of the Argentine Confederation. However, on the aftermath of 1852 Battle of Caseros, the Province of Buenos Aires had seceded from the Confederation. In 1859, after the Battle of Cepeda the State of Buenos Aires rejoined the Confederation, although it was granted the right to make some amendments to its Constitution. Finally, after the 1861 Battle of Pavón, Buenos Aires took over the Confederation.

The following federal governments fought the weaker Federalist and Autonomist resistances in the countryside until the 1870s. The last Autonomist rebellion in Buenos Aires was quelled in 1880, leading to the federalization of Buenos Aires city and the stabilization of the Argentine State and government through the National Autonomist Party.

Quebec
Federalism, in regard to the National Question, refers to support for Quebec remaining within Canada, while either keeping the status quo or pursuing greater autonomy and constitutional recognition of a Quebec nation, with corresponding rights and powers for Quebec within the Canadian federation. This ideology is opposed to Quebec sovereigntism, proponents of Quebec independence, most often (but not for all followers) along with an economic union with Canada similar to the European Union.

United States
In the United States the term federalist usually applies to a member of one of the following groups:
 * Statesmen and public figures supporting the proposed Constitution of the United States between 1787 and 1789. The most prominent advocates were James Madison, Alexander Hamilton, and John Jay. They published The Federalist Papers, which expounded the principles of the early federalist movement to promote and adopt the proposed Constitution.
 * Statesmen and public figures supporting the administrations of presidents George Washington (1789–1797) and John Adams (1797–1801). They became the Federalist Party, founded by Alexander Hamilton. During the 1790s and early 1800s, the Federalist Party opposed the Democratic-Republican Party (founded by Thomas Jefferson and James Madison) over issues of how broadly or narrowly to apply the provisions of the new Constitution.

The Federalist Society for Law and Public Policy Studies is an organization of conservative and libertarian lawyers and others dedicated to debate of these principles.

Global federalism
The World Federalist Movement is a global citizens movement that advocates for strengthened and democratic world institutions subjected to the federalist principles of subsidiarity, solidarity and democracy. It states that "[w]orld federalists support the creation of democratic global structures accountable to the citizens of the world and call for the division of international authority among separate agencies".

title: Federalist