title: Mitochondrion

text: Such trafficking capacity depends on the MAM, which has been shown to facilitate transfer of lipid intermediates between organelles. In contrast to the standard vesicular mechanism of lipid transfer, evidence indicates that the physical proximity of the ER and mitochondrial membranes at the MAM allows for lipid flipping between opposed bilayers. Despite this unusual and seemingly energetically unfavorable mechanism, such transport does not require ATP. Instead, in yeast, it has been shown to be dependent on a multiprotein tethering structure termed the ER-mitochondria encounter structure, or ERMES, although it remains unclear whether this structure directly mediates lipid transfer or is required to keep the membranes in sufficiently close proximity to lower the energy barrier for lipid flipping.


title: Mitochondrion

text: The MAM may also be part of the secretory pathway, in addition to its role in intracellular lipid trafficking. In particular, the MAM appears to be an intermediate destination between the rough ER and the Golgi in the pathway that leads to very-low-density lipoprotein, or VLDL, assembly and secretion. The MAM thus serves as a critical metabolic and trafficking hub in lipid metabolism.


title: Mitochondrion

text: A critical role for the ER in calcium signaling was acknowledged before such a role for the mitochondria was widely accepted, in part because the low affinity of Ca channels localized to the outer mitochondrial membrane seemed to contradict this organelle's purported responsiveness to changes in intracellular Ca flux. But the presence of the MAM resolves this apparent contradiction: the close physical association between the two organelles results in Ca microdomains at contact points that facilitate efficient Ca transmission from the ER to the mitochondria. Transmission occurs in response to so-called "Ca puffs" generated by spontaneous clustering and activation of IP3R, a canonical ER membrane Ca channel.


title: Mitochondrion

text: The fate of these puffs—in particular, whether they remain restricted to isolated locales or integrated into Ca waves for propagation throughout the cell—is determined in large part by MAM dynamics. Although reuptake of Ca by the ER (concomitant with its release) modulates the intensity of the puffs, thus insulating mitochondria to a certain degree from high Ca exposure, the MAM often serves as a firewall that essentially buffers Ca puffs by acting as a sink into which free ions released into the cytosol can be funneled. This Ca tunneling occurs through the low-affinity Ca receptor VDAC1, which recently has been shown to be physically tethered to the IP3R clusters on the ER membrane and enriched at the MAM. The ability of mitochondria to serve as a Ca sink is a result of the electrochemical gradient generated during oxidative phosphorylation, which makes tunneling of the cation an exergonic process. Normal, mild calcium influx from cytosol into the mitochondrial matrix causes transient depolarization that is corrected by pumping out protons.


title: Mitochondrion

text: But transmission of Ca is not unidirectional; rather, it is a two-way street. The properties of the Ca pump SERCA and the channel IP3R present on the ER membrane facilitate feedback regulation coordinated by MAM function. In particular, the clearance of Ca by the MAM allows for spatio-temporal patterning of Ca signaling because Ca alters IP3R activity in a biphasic manner. SERCA is likewise affected by mitochondrial feedback: uptake of Ca by the MAM stimulates ATP production, thus providing energy that enables SERCA to reload the ER with Ca for continued Ca efflux at the MAM. Thus, the MAM is not a passive buffer for Ca puffs; rather it helps modulate further Ca signaling through feedback loops that affect ER dynamics.


title: Mitochondrion

text: Regulating ER release of Ca at the MAM is especially critical because only a certain window of Ca uptake sustains the mitochondria, and consequently the cell, at homeostasis. Sufficient intraorganelle Ca signaling is required to stimulate metabolism by activating dehydrogenase enzymes critical to flux through the citric acid cycle. However, once Ca signaling in the mitochondria passes a certain threshold, it stimulates the intrinsic pathway of apoptosis in part by collapsing the mitochondrial membrane potential required for metabolism. Studies examining the role of pro- and anti-apoptotic factors support this model; for example, the anti-apoptotic factor Bcl-2 has been shown to interact with IP3Rs to reduce Ca filling of the ER, leading to reduced efflux at the MAM and preventing collapse of the mitochondrial membrane potential post-apoptotic stimuli. Given the need for such fine regulation of Ca signaling, it is perhaps unsurprising that dysregulated mitochondrial Ca has been implicated in several neurodegenerative diseases, while the catalogue of tumor suppressors includes a few that are enriched at the MAM.


title: Mitochondrion

text: Recent advances in the identification of the tethers between the mitochondrial and ER membranes suggest that the scaffolding function of the molecular elements involved is secondary to other, non-structural functions. In yeast, ERMES, a multiprotein complex of interacting ER- and mitochondrial-resident membrane proteins, is required for lipid transfer at the MAM and exemplifies this principle. One of its components, for example, is also a constituent of the protein complex required for insertion of transmembrane beta-barrel proteins into the lipid bilayer. However, a homologue of the ERMES complex has not yet been identified in mammalian cells. Other proteins implicated in scaffolding likewise have functions independent of structural tethering at the MAM; for example, ER-resident and mitochondrial-resident mitofusins form heterocomplexes that regulate the number of inter-organelle contact sites, although mitofusins were first identified for their role in fission and fusion events between individual mitochondria. Glucose-related protein 75 (grp75) is another dual-function protein. In addition to the matrix pool of grp75, a portion serves as a chaperone that physically links the mitochondrial and ER Ca channels VDAC and IP3R for efficient Ca transmission at the MAM. Another potential tether is Sigma-1R, a non-opioid receptor whose stabilization of ER-resident IP3R may preserve communication at the MAM during the metabolic stress response.


title: Mitochondrion

text: The MAM is a critical signaling, metabolic, and trafficking hub in the cell that allows for the integration of ER and mitochondrial physiology. Coupling between these organelles is not simply structural but functional as well and critical for overall cellular physiology and homeostasis. The MAM thus offers a perspective on mitochondria that diverges from the traditional view of this organelle as a static, isolated unit appropriated for its metabolic capacity by the cell. Instead, this mitochondrial-ER interface emphasizes the integration of the mitochondria, the product of an endosymbiotic event, into diverse cellular processes. Recently it has also been shown, that mitochondria and MAM-s in neurons are anchored to specialised intercellular communication sites (so called somatic-junctions). Microglial processes monitor and protect neuronal functions at these sites, and MAM-s are supposed to have an important role in this type of cellular quality-control.


title: Mitochondrion

text: There are two hypotheses about the origin of mitochondria: endosymbiotic and autogenous. The endosymbiotic hypothesis suggests that mitochondria were originally prokaryotic cells, capable of implementing oxidative mechanisms that were not possible for eukaryotic cells; they became endosymbionts living inside the eukaryote. In the autogenous hypothesis, mitochondria were born by splitting off a portion of DNA from the nucleus of the eukaryotic cell at the time of divergence with the prokaryotes; this DNA portion would have been enclosed by membranes, which could not be crossed by proteins. Since mitochondria have many features in common with bacteria, the endosymbiotic hypothesis is the more widely accepted of the two accounts.


title: Mitochondrion

text: A mitochondrion contains DNA, which is organized as several copies of a single, usually circular chromosome. This mitochondrial chromosome contains genes for redox proteins, such as those of the respiratory chain. The CoRR hypothesis proposes that this co-location is required for redox regulation. The mitochondrial genome codes for some RNAs of ribosomes, and the 22 tRNAs necessary for the translation of mRNAs into protein. The circular structure is also found in prokaryotes. The proto-mitochondrion was probably closely related to "Rickettsia". However, the exact relationship of the ancestor of mitochondria to the alphaproteobacteria and whether the mitochondrion was formed at the same time or after the nucleus, remains controversial. For example, it has been suggested that the SAR11 clade of bacteria shares a relatively recent common ancestor with the mitochondria, while phylogenomic analyses indicate that mitochondria evolved from a Pseudomonadota lineage that is closely related to or a member of alphaproteobacteria. Some papers describe mitochondria as sister to the alphaproteobactera, together forming the sister the marineproteo1 group, together forming the sister to Magnetococcidae.


title: Mitochondrion

text: The ribosomes coded for by the mitochondrial DNA are similar to those from bacteria in size and structure. They closely resemble the bacterial 70S ribosome and not the 80S cytoplasmic ribosomes, which are coded for by nuclear DNA.


title: Mitochondrion

text: The endosymbiotic relationship of mitochondria with their host cells was popularized by Lynn Margulis. The endosymbiotic hypothesis suggests that mitochondria descended from aerobic bacteria that somehow survived endocytosis by another cell, and became incorporated into the cytoplasm. The ability of these bacteria to conduct respiration in host cells that had relied on glycolysis and fermentation would have provided a considerable evolutionary advantage. This symbiotic relationship probably developed 1.7 to 2 billion years ago.


title: Mitochondrion

text: A few groups of unicellular eukaryotes have only vestigial mitochondria or derived structures: The microsporidians, metamonads, and archamoebae. These groups appear as the most primitive eukaryotes on phylogenetic trees constructed using rRNA information, which once suggested that they appeared before the origin of mitochondria. However, this is now known to be an artifact of "long-branch attraction": They are derived groups and retain genes or organelles derived from mitochondria (e. g., mitosomes and hydrogenosomes). Hydrogenosomes, mitosomes, and related organelles as found in some loricifera (e. g. "Spinoloricus") and myxozoa (e. g. "Henneguya zschokkei") are together classified as MROs, mitochondrion-related organelles.


title: Mitochondrion

text: Monocercomonoides appear to have lost their mitochondria completely and at least some of the mitochondrial functions seem to be carried out by cytoplasmic proteins now"."


title: Mitochondrion

text: Mitochondria contain their own genome. The human mitochondrial genome is a circular double-stranded DNA molecule of about 16 kilobases. It encodes 37 genes: 13 for subunits of respiratory complexes I, III, IV and V, 22 for mitochondrial tRNA (for the 20 standard amino acids, plus an extra gene for leucine and serine), and 2 for rRNA (12S and 16S rRNA). One mitochondrion can contain two to ten copies of its DNA. One of the two mitochondrial DNA (mtDNA) strands has a disproportionately higher ratio of the heavier nucleotides adenine and guanine, and this is termed the heavy strand (or H strand), whereas the other strand is termed the light strand (or L strand). The weight difference allows the two strands to be separated by centrifugation. mtDNA has one long non-coding stretch known as the non-coding region (NCR), which contains the heavy strand promoter (HSP) and light strand promoter (LSP) for RNA transcription, the origin of replication for the H strand (OriH) localized on the L strand, three conserved sequence boxes (CSBs 1–3), and a termination-associated sequence (TAS). The origin of replication for the L strand (OriL) is localized on the H strand 11,000 bp downstream of OriH, located within a cluster of genes coding for tRNA.


title: Mitochondrion

text: As in prokaryotes, there is a very high proportion of coding DNA and an absence of repeats. Mitochondrial genes are transcribed as multigenic transcripts, which are cleaved and polyadenylated to yield mature mRNAs. Most proteins necessary for mitochondrial function are encoded by genes in the cell nucleus and the corresponding proteins are imported into the mitochondrion. The exact number of genes encoded by the nucleus and the mitochondrial genome differs between species. Most mitochondrial genomes are circular. In general, mitochondrial DNA lacks introns, as is the case in the human mitochondrial genome; however, introns have been observed in some eukaryotic mitochondrial DNA, such as that of yeast and protists, including "Dictyostelium discoideum". Between protein-coding regions, tRNAs are present. Mitochondrial tRNA genes have different sequences from the nuclear tRNAs, but lookalikes of mitochondrial tRNAs have been found in the nuclear chromosomes with high sequence similarity.


title: Mitochondrion

text: In animals, the mitochondrial genome is typically a single circular chromosome that is approximately 16 kb long and has 37 genes. The genes, while highly conserved, may vary in location. Curiously, this pattern is not found in the human body louse ("Pediculus humanus"). Instead, this mitochondrial genome is arranged in 18 minicircular chromosomes, each of which is 3–4 kb long and has one to three genes. This pattern is also found in other sucking lice, but not in chewing lice. Recombination has been shown to occur between the minichromosomes.


title: Mitochondrion

text: The near-absence of genetic recombination in mitochondrial DNA makes it a useful source of information for studying population genetics and evolutionary biology. Because all the mitochondrial DNA is inherited as a single unit, or haplotype, the relationships between mitochondrial DNA from different individuals can be represented as a gene tree. Patterns in these gene trees can be used to infer the evolutionary history of populations. The classic example of this is in human evolutionary genetics, where the molecular clock can be used to provide a recent date for mitochondrial Eve. This is often interpreted as strong support for a recent modern human expansion out of Africa. Another human example is the sequencing of mitochondrial DNA from Neanderthal bones. The relatively large evolutionary distance between the mitochondrial DNA sequences of Neanderthals and living humans has been interpreted as evidence for the lack of interbreeding between Neanderthals and modern humans.


title: Mitochondrion

text: However, mitochondrial DNA reflects only the history of the females in a population. This can be partially overcome by the use of paternal genetic sequences, such as the non-recombining region of the Y-chromosome.


title: Mitochondrion

text: Recent measurements of the molecular clock for mitochondrial DNA reported a value of 1 mutation every 7884 years dating back to the most recent common ancestor of humans and apes, which is consistent with estimates of mutation rates of autosomal DNA (10 per base per generation).


title: Mitochondrion

text: While slight variations on the standard genetic code had been predicted earlier, none was discovered until 1979, when researchers studying human mitochondrial genes determined that they used an alternative code. Nonetheless, the mitochondria of many other eukaryotes, including most plants, use the standard code. Many slight variants have been discovered since, including various alternative mitochondrial codes. Further, the AUA, AUC, and AUU codons are all allowable start codons.


title: Mitochondrion

text: Some of these differences should be regarded as pseudo-changes in the genetic code due to the phenomenon of RNA editing, which is common in mitochondria. In higher plants, it was thought that CGG encoded for tryptophan and not arginine; however, the codon in the processed RNA was discovered to be the UGG codon, consistent with the standard genetic code for tryptophan. Of note, the arthropod mitochondrial genetic code has undergone parallel evolution within a phylum, with some organisms uniquely translating AGG to lysine.


title: Mitochondrion

text: Mitochondria divide by mitochondrial fission, a form of binary fission that is also done by bacteria although the process is tightly regulated by the host eukaryotic cell and involves communication between and contact with several other organelles. The regulation of this division differs between eukaryotes. In many single-celled eukaryotes, their growth and division are linked to the cell cycle. For example, a single mitochondrion may divide synchronously with the nucleus. This division and segregation process must be tightly controlled so that each daughter cell receives at least one mitochondrion. In other eukaryotes (in mammals for example), mitochondria may replicate their DNA and divide mainly in response to the energy needs of the cell, rather than in phase with the cell cycle. When the energy needs of a cell are high, mitochondria grow and divide. When energy use is low, mitochondria are destroyed or become inactive. In such examples mitochondria are apparently randomly distributed to the daughter cells during the division of the cytoplasm. Mitochondrial dynamics, the balance between mitochondrial fusion and fission, is an important factor in pathologies associated with several disease conditions.


title: Mitochondrion

text: The hypothesis of mitochondrial binary fission has relied on the visualization by fluorescence microscopy and conventional transmission electron microscopy (TEM). The resolution of fluorescence microscopy (~200 nm) is insufficient to distinguish structural details, such as double mitochondrial membrane in mitochondrial division or even to distinguish individual mitochondria when several are close together. Conventional TEM has also some technical limitations in verifying mitochondrial division. Cryo-electron tomography was recently used to visualize mitochondrial division in frozen hydrated intact cells. It revealed that mitochondria divide by budding.


title: Mitochondrion

text: An individual's mitochondrial genes are inherited only from the mother, with rare exceptions. In humans, when an egg cell is fertilized by a sperm, the mitochondria, and therefore the mitochondrial DNA, usually come from the egg only. The sperm's mitochondria enter the egg, but do not contribute genetic information to the embryo. Instead, paternal mitochondria are marked with ubiquitin to select them for later destruction inside the embryo. The egg cell contains relatively few mitochondria, but these mitochondria divide to populate the cells of the adult organism. This mode is seen in most organisms, including the majority of animals. However, mitochondria in some species can sometimes be inherited paternally. This is the norm among certain coniferous plants, although not in pine trees and yews. For Mytilids, paternal inheritance only occurs within males of the species. It has been suggested that it occurs at a very low level in humans.


title: Mitochondrion

text: Uniparental inheritance leads to little opportunity for genetic recombination between different lineages of mitochondria, although a single mitochondrion can contain 2–10 copies of its DNA. What recombination does take place maintains genetic integrity rather than maintaining diversity. However, there are studies showing evidence of recombination in mitochondrial DNA. It is clear that the enzymes necessary for recombination are present in mammalian cells. Further, evidence suggests that animal mitochondria can undergo recombination. The data are more controversial in humans, although indirect evidence of recombination exists.


title: Mitochondrion

text: Entities undergoing uniparental inheritance and with little to no recombination may be expected to be subject to Muller's ratchet, the accumulation of deleterious mutations until functionality is lost. Animal populations of mitochondria avoid this buildup through a developmental process known as the mtDNA bottleneck. The bottleneck exploits stochastic processes in the cell to increase the cell-to-cell variability in mutant load as an organism develops: a single egg cell with some proportion of mutant mtDNA thus produces an embryo where different cells have different mutant loads. Cell-level selection may then act to remove those cells with more mutant mtDNA, leading to a stabilization or reduction in mutant load between generations. The mechanism underlying the bottleneck is debated, with a recent mathematical and experimental metastudy providing evidence for a combination of random partitioning of mtDNAs at cell divisions and random turnover of mtDNA molecules within the cell.


title: Mitochondrion

text: Mitochondria can repair oxidative DNA damage by mechanisms analogous to those occurring in the cell nucleus. The proteins employed in mtDNA repair are encoded by nuclear genes, and are translocated to the mitochondria. The DNA repair pathways in mammalian mitochondria include base excision repair, double-strand break repair, direct reversal and mismatch repair. Alternatively, DNA damage may be bypassed, rather than repaired, by translesion synthesis.


title: Mitochondrion

text: Of the several DNA repair process in mitochondria, the base excision repair pathway has been most comprehensively studied. Base excision repair is carried out by a sequence of enzyme-catalyzed steps that include recognition and excision of a damaged DNA base, removal of the resulting abasic site, end processing, gap filling and ligation. A common damage in mtDNA that is repaired by base excision repair is 8-oxoguanine produced by oxidation of guanine.


title: Mitochondrion

text: Double-strand breaks can be repaired by homologous recombinational repair in both mammalian mtDNA and plant mtDNA. Double-strand breaks in mtDNA can also be repaired by microhomology-mediated end joining. Although there is evidence for the repair processes of direct reversal and mismatch repair in mtDNA, these processes are not well characterized.


title: Mitochondrion

text: Some organisms have lost mitochondrial DNA altogether. In these cases, genes encoded by the mitochondrial DNA have been lost or transferred to the nucleus. "Cryptosporidium" have mitochondria that lack any DNA, presumably because all their genes have been lost or transferred. In "Cryptosporidium", the mitochondria have an altered ATP generation system that renders the parasite resistant to many classical mitochondrial inhibitors such as cyanide, azide, and atovaquone. Mitochondria that lack their own DNA have been found in a marine parasitic dinoflagellate from the genus "Amoebophyra". This microorganism, "A. cerati", has functional mitochondria that lack a genome. In related species, the mitochondrial genome still has three genes, but in "A. cerati" only a single mitochondrial gene — the cytochrome c oxidase I gene ("cox1") — is found, and it has migrated to the genome of the nucleus.


title: Mitochondrion

text: Damage and subsequent dysfunction in mitochondria is an important factor in a range of human diseases due to their influence in cell metabolism. Mitochondrial disorders often present as neurological disorders, including autism. They can also manifest as myopathy, diabetes, multiple endocrinopathy, and a variety of other systemic disorders. Diseases caused by mutation in the mtDNA include Kearns–Sayre syndrome, MELAS syndrome and Leber's hereditary optic neuropathy. In the vast majority of cases, these diseases are transmitted by a female to her children, as the zygote derives its mitochondria and hence its mtDNA from the ovum. Diseases such as Kearns-Sayre syndrome, Pearson syndrome, and progressive external ophthalmoplegia are thought to be due to large-scale mtDNA rearrangements, whereas other diseases such as MELAS syndrome, Leber's hereditary optic neuropathy, MERRF syndrome, and others are due to point mutations in mtDNA.


title: Mitochondrion

text: It has also been reported that drug tolerant cancer cells have an increased number and size of mitochondria which suggested an increase in mitochondrial biogenesis. Interestingly, a recent study in "Nature Nanotechnology" has reported that cancer cells can hijack the mitochondria from immune cells via physical tunneling nanotubes.


title: Mitochondrion

text: In other diseases, defects in nuclear genes lead to dysfunction of mitochondrial proteins. This is the case in Friedreich's ataxia, hereditary spastic paraplegia, and Wilson's disease. These diseases are inherited in a dominance relationship, as applies to most other genetic diseases. A variety of disorders can be caused by nuclear mutations of oxidative phosphorylation enzymes, such as coenzyme Q10 deficiency and Barth syndrome. Environmental influences may interact with hereditary predispositions and cause mitochondrial disease. For example, there may be a link between pesticide exposure and the later onset of Parkinson's disease. Other pathologies with etiology involving mitochondrial dysfunction include schizophrenia, bipolar disorder, dementia, Alzheimer's disease, Parkinson's disease, epilepsy, stroke, cardiovascular disease, chronic fatigue syndrome, retinitis pigmentosa, and diabetes mellitus.


title: Mitochondrion

text: Mitochondria-mediated oxidative stress plays a role in cardiomyopathy in type 2 diabetics. Increased fatty acid delivery to the heart increases fatty acid uptake by cardiomyocytes, resulting in increased fatty acid oxidation in these cells. This process increases the reducing equivalents available to the electron transport chain of the mitochondria, ultimately increasing reactive oxygen species (ROS) production. ROS increases uncoupling proteins (UCPs) and potentiate proton leakage through the adenine nucleotide translocator (ANT), the combination of which uncouples the mitochondria. Uncoupling then increases oxygen consumption by the mitochondria, compounding the increase in fatty acid oxidation. This creates a vicious cycle of uncoupling; furthermore, even though oxygen consumption increases, ATP synthesis does not increase proportionally because the mitochondria are uncoupled. Less ATP availability ultimately results in an energy deficit presenting as reduced cardiac efficiency and contractile dysfunction. To compound the problem, impaired sarcoplasmic reticulum calcium release and reduced mitochondrial reuptake limits peak cytosolic levels of the important signaling ion during muscle contraction. Decreased intra-mitochondrial calcium concentration increases dehydrogenase activation and ATP synthesis. So in addition to lower ATP synthesis due to fatty acid oxidation, ATP synthesis is impaired by poor calcium signaling as well, causing cardiac problems for diabetics.


title: Mitochondrion

text: There may be some leakage of the electrons transferred in the respiratory chain to form reactive oxygen species. This was thought to result in significant oxidative stress in the mitochondria with high mutation rates of mitochondrial DNA. Hypothesized links between aging and oxidative stress are not new and were proposed in 1956, which was later refined into the mitochondrial free radical theory of aging. A vicious cycle was thought to occur, as oxidative stress leads to mitochondrial DNA mutations, which can lead to enzymatic abnormalities and further oxidative stress.


title: Mitochondrion

text: A number of changes can occur to mitochondria during the aging process. Tissues from elderly humans show a decrease in enzymatic activity of the proteins of the respiratory chain. However, mutated mtDNA can only be found in about 0.2% of very old cells. Large deletions in the mitochondrial genome have been hypothesized to lead to high levels of oxidative stress and neuronal death in Parkinson's disease. Mitochondrial dysfunction has also been shown to occur in amyotrophic lateral sclerosis.


title: Mitochondrion

text: Since mitochondria cover a pivotal role in the ovarian function, by providing ATP necessary for the development from germinal vesicle to mature oocyte, a decreased mitochondria function can lead to inflammation, resulting in premature ovarian failure and accelerated ovarian aging. The resulting dysfunction is then reflected in quantitative (such as mtDNA copy number and mtDNA deletions), qualitative (such as mutations and strand breaks) and oxidative damage (such as dysfunctional mitochondria due to ROS), which are not only relevant in ovarian aging, but perturb oocyte-cumulus crosstalk in the ovary, are linked to genetic disorders (such as Fragile X) and can interfere with embryo selection.


title: Mitochondrion

text: The first observations of intracellular structures that probably represented mitochondria were published in 1857, by the physiologist Albert von Kolliker. Richard Altmann, in 1890, established them as cell organelles and called them "bioblasts." In 1898, Carl Benda coined the term "mitochondria" from the Greek , , "thread", and , , "granule." Leonor Michaelis discovered that Janus green can be used as a supravital stain for mitochondria in 1900. In 1904, Friedrich Meves, made the first recorded observation of mitochondria in plants in cells of the white waterlily, "Nymphaea alba" and in 1908, along with Claudius Regaud, suggested that they contain proteins and lipids. Benjamin F. Kingsbury, in 1912, first related them with cell respiration, but almost exclusively based on morphological observations. In 1913, particles from extracts of guinea-pig liver were linked to respiration by Otto Heinrich Warburg, which he called "grana." Warburg and Heinrich Otto Wieland, who had also postulated a similar particle mechanism, disagreed on the chemical nature of the respiration. It was not until 1925, when David Keilin discovered cytochromes, that the respiratory chain was described.


title: Mitochondrion

text: In 1939, experiments using minced muscle cells demonstrated that cellular respiration using one oxygen molecule can form four adenosine triphosphate (ATP) molecules, and in 1941, the concept of the phosphate bonds of ATP being a form of energy in cellular metabolism was developed by Fritz Albert Lipmann. In the following years, the mechanism behind cellular respiration was further elaborated, although its link to the mitochondria was not known. The introduction of tissue fractionation by Albert Claude allowed mitochondria to be isolated from other cell fractions and biochemical analysis to be conducted on them alone. In 1946, he concluded that cytochrome oxidase and other enzymes responsible for the respiratory chain were isolated to the mitochondria. Eugene Kennedy and Albert Lehninger discovered in 1948 that mitochondria are the site of oxidative phosphorylation in eukaryotes. Over time, the fractionation method was further developed, improving the quality of the mitochondria isolated, and other elements of cell respiration were determined to occur in the mitochondria.


title: Mitochondrion

text: The first high-resolution electron micrographs appeared in 1952, replacing the Janus Green stains as the preferred way to visualize mitochondria. This led to a more detailed analysis of the structure of the mitochondria, including confirmation that they were surrounded by a membrane. It also showed a second membrane inside the mitochondria that folded up in ridges dividing up the inner chamber and that the size and shape of the mitochondria varied from cell to cell.


title: Mitochondrion

text: In 1967, it was discovered that mitochondria contained ribosomes. In 1968, methods were developed for mapping the mitochondrial genes, with the genetic and physical map of yeast mitochondrial DNA completed in 1976.


title: Gunpowder Plot

text: The Gunpowder Plot of 1605, in earlier centuries often called the Gunpowder Treason Plot or the Jesuit Treason, was a failed assassination attempt against King James I by a group of provincial English Catholics led by Robert Catesby who sought to restore the Catholic monarchy to England after decades of persecution against Catholics.


title: Gunpowder Plot

text: The plan was to blow up the House of Lords during the State Opening of Parliament on 5 November 1605, as the prelude to a popular revolt in the Midlands during which King James's nine-year-old daughter, Elizabeth, was to be installed as the Catholic head of state. Catesby may have embarked on the scheme after hopes of securing greater religious tolerance under King James I had faded, leaving many English Catholics disappointed. His fellow contributors were John and Christopher Wright, Robert and Thomas Wintour, Thomas Percy, Guy Fawkes, Robert Keyes, Thomas Bates, John Grant, Ambrose Rookwood, Sir Everard Digby and Francis Tresham. Fawkes, who had 10 years of military experience fighting in the Spanish Netherlands in the failed suppression of the Dutch Revolt, was given charge of the explosives.


title: Gunpowder Plot

text: The plot was revealed to the authorities in an anonymous letter sent to William Parker, 4th Baron Monteagle, on 26 October 1605. During a search of the House of Lords in the evening on 4 November 1605, Fawkes was discovered guarding 36 barrels of gunpowder—enough to reduce the House of Lords to rubble—and arrested. Most of the conspirators fled from London as they learned that the plot had been discovered, trying to enlist support along the way. Several made a stand against the pursuing Sheriff of Worcester and his men at Holbeche House; in the ensuing battle Catesby was one of those shot and killed. At their trial on 27 January 1606, eight of the survivors, including Fawkes, were convicted and sentenced to be hanged, drawn and quartered.


title: Gunpowder Plot

text: Details of the assassination attempt were allegedly known by the principal Jesuit of England, Father Henry Garnet . Although he was convicted of treason and sentenced to death, doubt has been cast on how much he really knew of the plot. As its existence was revealed to him through confession, Garnet was prevented from informing the authorities by the absolute confidentiality of the confessional. Although anti-Catholic legislation was introduced soon after the discovery of the plot, many important and loyal Catholics retained high office during King James I's reign. The thwarting of the Gunpowder Plot was commemorated for many years afterwards by special sermons and other public events such as the ringing of church bells, which evolved into the British variant of Bonfire Night of today.


title: Gunpowder Plot

text: Between 1533 and 1540, King Henry VIII took control of the English Church from Rome, the start of several decades of religious tension in England. English Catholics struggled in a society dominated by the newly separate and increasingly Protestant Church of England. Henry's daughter, Queen Elizabeth I, responded to the growing religious divide by introducing the Elizabethan Religious Settlement, which required anyone appointed to a public or church office to swear allegiance to the monarch as head of the Church and state. The penalties for refusal were severe; fines were imposed for recusancy, and repeat offenders risked imprisonment and execution. Catholicism became marginalised, but despite the threat of torture or execution, priests continued to practise their faith in secret.


title: Gunpowder Plot

text: Queen Elizabeth, unmarried and childless, steadfastly refused to name an heir. Many Catholics believed that her Catholic cousin, Mary, Queen of Scots, was the legitimate heir to the English throne, but she was executed for treason in 1587. The English Secretary of State, Robert Cecil, negotiated secretly with Mary's son and successor, King James VI of Scotland. In the months before Elizabeth's death on 24 March 1603, Cecil prepared the way for James to succeed her.


title: Gunpowder Plot

text: Some exiled Catholics favoured Philip II of Spain's daughter, Isabella, as Elizabeth's successor. More moderate Catholics looked to James's and Elizabeth's cousin Arbella Stuart, a woman thought to have Catholic sympathies. As Elizabeth's health deteriorated, the government detained those they considered to be the "principal papists", and the Privy Council grew so worried that Arbella Stuart was moved closer to London to prevent her from being kidnapped by papists.


title: Gunpowder Plot

text: Despite competing claims to the English throne, the transition of power following Elizabeth's death went smoothly. James's succession was announced by a proclamation from Cecil on 24 March, which was generally celebrated. Leading papists, rather than causing trouble as anticipated, reacted to the news by offering their enthusiastic support for the new monarch. Jesuit priests, whose presence in England was punishable by death, also demonstrated their support for James, who was widely believed to embody "the natural order of things".


title: Gunpowder Plot

text: James ordered a ceasefire in the conflict with Spain, and even though the two countries were still technically at war, King Philip III sent his envoy, Don Juan de Tassis, to congratulate James on his accession. In the following year both countries signed the Treaty of London.


title: Gunpowder Plot

text: For decades, the English had lived under a monarch who refused to provide an heir, but James arrived with a family and a clear line of succession. His wife, Anne of Denmark, was the daughter of King Frederick II of Denmark. Their eldest child, the nine-year-old Henry, was considered a handsome and confident boy, and their two younger children, Elizabeth and Charles, were proof that James was able to provide heirs to continue the Protestant monarchy.


title: Gunpowder Plot

text: James's attitude towards Catholics was more moderate than that of his predecessor, perhaps even tolerant. He swore that he would not "persecute any that will be quiet and give an outward obedience to the law", and believed that exile was a better solution than capital punishment: "I would be glad to have both their heads and their bodies separated from this whole island and transported beyond seas." Some Catholics believed that the martyrdom of James's mother, Mary, Queen of Scots, would encourage James to convert to the Catholic faith, and the Catholic houses of Europe may also have shared that hope.


title: Gunpowder Plot

text: James received an envoy from Albert VII, ruler of the remaining Catholic territories in the Netherlands after over 30 years of war in the Dutch Revolt by English-supported Protestant rebels. For the Catholic expatriates engaged in that struggle, the restoration by force of a Catholic monarchy was an intriguing possibility, but following the failed Spanish invasion of England in 1588 the papacy had taken a longer-term view on the return of a Catholic monarch to the English throne.


title: Gunpowder Plot

text: During James I's reign the European wars of religion were intensifying. Protestants and Catholics were engaged in violent persecution of each other across Europe following the Protestant Reformation. Catholics made several assassination attempts on Protestant rulers in Europe and in England, including plans to poison James I's predecessor, Elizabeth I. In 1589, during the French Wars of Religion, the French King Henry III was mortally wounded with a dagger by Jacques Clément, a fanatic member of the Catholic League of France. Nine years later, the Jesuit Juan de Mariana's 1599 "On Kings and the Education of Kings" ("De rege et regis institutione") argued in support of tyrannicide. This work recounted the assassination of Henry III and argued for the legal right to overthrow a tyrant. Perhaps due in part to the publication of "De rege", until the 1620s, some English Catholics believed that regicide was justifiable to remove 'tyrants' from power. Much of the "rather nervous" political writing from James I was "concerned with the threat of Catholic assassination and refutation of the [Catholic] argument that 'faith did not need to be kept with heretics'".


title: Gunpowder Plot

text: In the absence of any sign that James would move to end the persecution of Catholics, as some had hoped for, several members of the clergy (including two anti-Jesuit priests) decided to take matters into their own hands. In what became known as the Bye Plot, the priests William Watson and William Clark planned to kidnap James and hold him in the Tower of London until he agreed to be more tolerant towards Catholics. Cecil received news of the plot from several sources, including the Archpriest George Blackwell, who instructed his priests to have no part in any such schemes. At about the same time, Lord Cobham, Lord Grey de Wilton, Griffin Markham and Walter Raleigh hatched what became known as the Main Plot, which involved removing James and his family and supplanting them with Arbella Stuart. Amongst others, they approached Philip III of Spain for funding, but were unsuccessful. All those involved in both plots were arrested in July and tried in autumn 1603. George Brooke was executed, but James—keen not to have too bloody a start to his reign—reprieved Cobham, Grey, and Markham while they were at the scaffold. Raleigh, who had watched while his colleagues sweated, had been due to be executed a few days later, but was also pardoned. Arbella Stuart denied any knowledge of the Main Plot. However, the two priests, Watson and Clark—condemned and "very bloodily handled"—were executed.


title: Gunpowder Plot

text: The Catholic community responded to news of these plots with shock. That the Bye Plot had been revealed by Catholics was instrumental in saving them from further persecution, and James was grateful enough to allow pardons for those recusants who sued for them, as well as postponing payment of their fines for a year.


title: Gunpowder Plot

text: On 19 February 1604, shortly after he discovered that his wife, Queen Anne, had been sent a rosary from the pope via one of James's spies, Sir Anthony Standen, James denounced the Catholic Church. Three days later, he ordered all Jesuits and all other Catholic priests to leave the country, and reimposed the collection of fines for recusancy.


title: Gunpowder Plot

text: James changed his focus from the anxieties of English Catholics to the establishment of an Anglo-Scottish union. He also appointed Scottish nobles such as George Home to his court, which proved unpopular with the Parliament of England. Some Members of Parliament made it clear that, in their view, the "effluxion of people from the Northern parts" was unwelcome, and compared them to "plants which are transported from barren ground into a more fertile one". Even more discontent resulted when the King allowed his Scottish nobles to collect the recusancy fines. There were 5,560 convicted of recusancy in 1605, of whom 112 were landowners. The very few Catholics of great wealth who refused to attend services at their parish church were fined £20 per month. Those of more moderate means had to pay two-thirds of their annual rental income; middle class recusants were fined one shilling a week, although the collection of all these fines was "haphazard and negligent". When James came to power, almost £5,000 a year (equivalent to almost £12 million in 2020) was being raised by these fines.


title: Gunpowder Plot

text: On 19 March, the King gave his opening speech to his first English Parliament in which he spoke of his desire to secure peace, but only by "profession of the true religion". He also spoke of a Christian union and reiterated his desire to avoid religious persecution. For the Catholics, the King's speech made it clear that they were not to "increase their number and strength in this Kingdom", that "they might be in hope to erect their Religion again". To Father John Gerard, these words were almost certainly responsible for the heightened levels of persecution the members of his faith now suffered, and for the priest Oswald Tesimond, they were a repudiation of the early claims that the King had made, upon which the papists had built their hopes. A week after James's speech, Lord Edmund Sheffield informed the king of over 900 recusants brought before the Assizes in Normanby, and on 24 April, the Popish Recusants Act 1605 was introduced in Parliament which threatened to outlaw all English followers of the Catholic Church.


title: Gunpowder Plot

text: The conspirators' principal aim was to kill King James, but many other important targets would also be present at the State Opening of Parliament, including the monarch's nearest relatives and members of the Privy Council. The senior judges of the English legal system, most of the Protestant aristocracy, and the bishops of the Church of England would all have attended in their capacity as members of the House of Lords, along with the members of the House of Commons. Another important objective was the kidnapping of the King's daughter, Elizabeth. Housed at Coombe Abbey near Coventry, she lived only ten miles north of Warwick—convenient for the plotters, most of whom lived in the Midlands. Once the King and his Parliament were dead, the plotters intended to install Elizabeth on the English throne as a titular Queen. The fate of her brothers, Henry and Charles, would be improvised; their role in state ceremonies was, as yet, uncertain. The plotters planned to use Henry Percy, 9th Earl of Northumberland, as Elizabeth's regent, but most likely never informed him of this.


title: Gunpowder Plot

text: Robert Catesby (1573–1605), a man of "ancient, historic and distinguished lineage", was the inspiration behind the plot. He was described by contemporaries as "a good-looking man, about six feet tall, athletic and a good swordsman". Along with several other conspirators, he took part in the Essex Rebellion in 1601, during which he was wounded and captured. Queen Elizabeth allowed him to escape with his life after fining him 4,000 marks (equivalent to more than £6 million in 2008), after which he sold his estate in Chastleton.


title: Gunpowder Plot

text: In 1603, Catesby helped to organise a mission to the new king of Spain, Philip III, urging Philip to launch an invasion attempt on England, which they assured him would be well supported, particularly by the English Catholics. Thomas Wintour (1571–1606) was chosen as the emissary, but the Spanish king, although sympathetic to the plight of Catholics in England, was intent on making peace with James. Wintour had also attempted to convince the Spanish envoy Don Juan de Tassis that "3,000 Catholics" were ready and waiting to support such an invasion. Concern was voiced by Pope Clement VIII that using violence to achieve a restoration of Catholic power in England would result in the destruction of those that remained.


title: Gunpowder Plot

text: According to contemporary accounts, in February 1604, Catesby invited Thomas Wintour to his house in Lambeth, where they discussed Catesby's plan to re-establish Catholicism in England by blowing up the House of Lords during the State Opening of Parliament. Wintour was known as a competent scholar, able to speak several languages, and he had fought with the English army in the Netherlands. His uncle, Francis Ingleby, had been executed for being a Catholic priest in 1586, and Wintour later converted to Catholicism. Also present at the meeting was John Wright, a devout Catholic said to be one of the best swordsmen of his day, and a man who had taken part with Catesby in the Earl of Essex's rebellion three years earlier. Despite his reservations over the possible repercussions should the attempt fail, Wintour agreed to join the conspiracy, perhaps persuaded by Catesby's rhetoric: "Let us give the attempt and where it faileth, pass no further."


title: Gunpowder Plot

text: Wintour travelled to Flanders to enquire about Spanish support. While there, he sought out Guy Fawkes (1570–1606), a committed Catholic who had served as a soldier in the Southern Netherlands under the command of William Stanley, and in 1603 had been recommended for a captaincy. Accompanied by John Wright's brother Christopher, Fawkes had also been a member of the 1603 delegation to the Spanish court pleading for an invasion of England. Wintour told Fawkes that "", and that certain gentlemen "". The two men returned to England late in April 1604, telling Catesby that Spanish support was unlikely. Thomas Percy, Catesby's friend and John Wright's brother-in-law, was introduced to the plot several weeks later.


title: Gunpowder Plot

text: Percy had found employment with his kinsman the Earl of Northumberland, and by 1596, was his agent for the family's northern estates. About 1600–1601 he served with his patron in the Low Countries. At some point during Northumberland's command in the Low Countries, Percy became his agent in his communications with James I. Percy was reputedly a "serious" character who had converted to the Catholic faith. His early years were, according to a Catholic source, marked by a tendency to rely on "his sword and personal courage". Northumberland, although not a Catholic himself, planned to build a strong relationship with James I in order to better the prospects of English Catholics, and to reduce the family disgrace caused by his separation from his wife Martha Wright, a favourite of Elizabeth I.


title: Gunpowder Plot

text: Thomas Percy's meetings with James seemed to go well. Percy returned with promises of support for the Catholics, and Northumberland believed that James would go so far as to allow Mass in private houses, so as not to cause public offence. Percy, keen to improve his standing, went even further, claiming that the future king would guarantee the safety of English Catholics.


title: Gunpowder Plot

text: The first meeting between the five conspirators took place on 20 May 1604, probably at the Duck and Drake Inn, just off the Strand, Thomas Wintour's usual residence when staying in London. Catesby, Thomas Wintour, and John Wright were in attendance, joined by Guy Fawkes and Thomas Percy. Alone in a private room, the five plotters swore an oath of secrecy on a prayer book. By coincidence, and ignorant of the plot, Father John Gerard (a friend of Catesby's) was celebrating Mass in another room, and the five men subsequently received the Eucharist.


title: Gunpowder Plot

text: Following their oath, the plotters left London and returned to their homes. The adjournment of Parliament gave them, they thought, until February 1605 to finalise their plans. On 9 June, Percy's patron, the Earl of Northumberland, appointed him to the Band of Gentlemen Pensioners, a mounted troop of 50 bodyguards to the King. This role gave Percy reason to seek a base in London, and a small property near the Prince's Chamber owned by Henry Ferrers, a tenant of John Whynniard, was chosen. Percy arranged for the use of the house through Northumberland's agents, Dudley Carleton and John Hippisley. Fawkes, using the pseudonym "John Johnson", took charge of the building, posing as Percy's servant.


title: Gunpowder Plot

text: The building was occupied by Scottish commissioners appointed by the King to consider his plans for the unification of England and Scotland, so the plotters hired Catesby's lodgings in Lambeth, on the opposite bank of the Thames, from where their stored gunpowder and other supplies could be conveniently rowed across each night. Meanwhile, King James I continued with his policies against the Catholics, and Parliament pushed through anti-Catholic legislation, until its adjournment on 7 July.


title: Gunpowder Plot

text: The conspirators returned to London in October 1604, when Robert Keyes, a "desperate man, ruined and indebted", was admitted to the group. His responsibility was to take charge of Catesby's house in Lambeth, where the gunpowder and other supplies were to be stored. Keyes's family had notable connections; his wife's employer was the Catholic Lord Mordaunt. He was tall, with a red beard, and was seen as trustworthy and—like Fawkes—capable of looking after himself. In December Catesby recruited his servant, Thomas Bates, into the plot, after the latter accidentally became aware of it.


title: Gunpowder Plot

text: It was announced on 24 December that the re-opening of Parliament would be delayed. Concern over the plague meant that rather than sitting in February, as the plotters had originally planned for, Parliament would not sit again until 3 October 1605. The contemporaneous account of the prosecution claimed that during this delay the conspirators were digging a tunnel beneath Parliament. This may have been a government fabrication, as no evidence for the existence of a tunnel was presented by the prosecution, and no trace of one has ever been found. The account of a tunnel comes directly from Thomas Wintour's confession, and Guy Fawkes did not admit the existence of such a scheme until his fifth interrogation. Logistically, digging a tunnel would have proved extremely difficult, especially as none of the conspirators had any experience of mining. If the story is true, by 6 December the Scottish commissioners had finished their work, and the conspirators were busy tunnelling from their rented house to the House of Lords. They ceased their efforts when, during tunnelling, they heard a noise from above. The noise turned out to be the then-tenant's widow, who was clearing out the undercroft directly beneath the House of Lords—the room where the plotters eventually stored the gunpowder.


title: Gunpowder Plot

text: By the time the plotters reconvened at the start of the old style new year on Lady Day, 25 March, three more had been admitted to their ranks; Robert Wintour, John Grant, and Christopher Wright. The additions of Wintour and Wright were obvious choices. Along with a small fortune, Robert Wintour inherited Huddington Court (a known refuge for priests) near Worcester, and was reputedly a generous and well-liked man. A devout Catholic, he married Gertrude, the daughter of John Talbot of Grafton, from a prominent Worcestershire family of recusants. Christopher Wright (1568–1605), John's brother, had also taken part in the Earl of Essex's revolt and had moved his family to Twigmore in Lincolnshire, then known as something of a haven for priests. John Grant was married to Wintour's sister, Dorothy, and was lord of the manor of Norbrook near Stratford-upon-Avon. Reputed to be an intelligent, thoughtful man, he sheltered Catholics at his home at Snitterfield, and was another who had been involved in the Essex revolt of 1601.


title: Gunpowder Plot

text: In addition, 25 March was the day on which the plotters purchased the lease to the undercroft they had supposedly tunnelled near to, owned by John Whynniard. The Palace of Westminster in the early 17th century was a warren of buildings clustered around the medieval chambers, chapels, and halls of the former royal palace that housed both Parliament and the various royal law courts. The old palace was easily accessible; merchants, lawyers, and others lived and worked in the lodgings, shops and taverns within its precincts. Whynniard's building was along a right-angle to the House of Lords, alongside a passageway called Parliament Place, which itself led to Parliament Stairs and the River Thames. Undercrofts were common features at the time, used to house a variety of materials including food and firewood. Whynniard's undercroft, on the ground floor, was directly beneath the first-floor House of Lords, and may once have been part of the palace's medieval kitchen. Unused and filthy, its location was ideal for what the group planned to do.


title: Gunpowder Plot

text: In the second week of June Catesby met in London the principal Jesuit in England, Father Henry Garnet, and asked him about the morality of entering into an undertaking which might involve the destruction of the innocent, together with the guilty. Garnet answered that such actions could often be excused, but according to his own account later admonished Catesby during a second meeting in July in Essex, showing him a letter from the pope which forbade rebellion. Soon after, the Jesuit priest Oswald Tesimond told Garnet he had taken Catesby's confession, in the course of which he had learnt of the plot. Garnet and Catesby met for a third time on 24 July 1605, at the house of the wealthy catholic Anne Vaux in Enfield Chase. Garnet decided that Tesimond's account had been given under the seal of the confessional, and that canon law therefore forbade him to repeat what he had heard. Without acknowledging that he was aware of the precise nature of the plot, Garnet attempted to dissuade Catesby from his course, to no avail. Garnet wrote to a colleague in Rome, Claudio Acquaviva, expressing his concerns about open rebellion in England. He also told Acquaviva that "there is a risk that some private endeavour may commit treason or use force against the King", and urged the pope to issue a public brief against the use of force.


title: Gunpowder Plot

text: According to Fawkes, 20 barrels of gunpowder were brought in at first, followed by 16 more on 20 July. The supply of gunpowder was theoretically controlled by the government, but it was easily obtained from illicit sources. On 28 July, the ever-present threat of the plague again delayed the opening of Parliament, this time until Tuesday 5 November. Fawkes left the country for a short time. The King, meanwhile, spent much of the summer away from the city, hunting. He stayed wherever was convenient, including on occasion at the houses of prominent Catholics. Garnet, convinced that the threat of an uprising had receded, travelled the country on a pilgrimage.


title: Gunpowder Plot

text: It is uncertain when Fawkes returned to England, but he was back in London by late August, when he and Wintour discovered that the gunpowder stored in the undercroft had decayed. More gunpowder was brought into the room, along with firewood to conceal it. The final three conspirators were recruited in late 1605. At Michaelmas, Catesby persuaded the staunchly Catholic Ambrose Rookwood to rent Clopton House near Stratford-upon-Avon. Rookwood was a young man with recusant connections, whose stable of horses at Coldham Hall in Stanningfield, Suffolk was an important factor in his enlistment. His parents, Robert Rookwood and Dorothea Drury, were wealthy landowners, and had educated their son at a Jesuit school near Calais. Everard Digby was a young man who was generally well liked, and lived at Gayhurst House in Buckinghamshire. He had been knighted by the King in April 1603, and was converted to Catholicism by Gerard. Digby and his wife, Mary Mulshaw, had accompanied the priest on his pilgrimage, and the two men were reportedly close friends. Digby was asked by Catesby to rent Coughton Court near Alcester. Digby also promised £1,500 after Percy failed to pay the rent due for the properties he had taken in Westminster. Finally, on 14 October Catesby invited Francis Tresham into the conspiracy. Tresham was the son of the Catholic Thomas Tresham, and a cousin to Robert Catesby; the two had been raised together. He was also the heir to his father's large fortune, which had been depleted by recusant fines, expensive tastes, and by Francis and Catesby's involvement in the Essex revolt.


title: Gunpowder Plot

text: Catesby and Tresham met at the home of Tresham's brother-in-law and cousin, Lord Stourton. In his confession, Tresham claimed that he had asked Catesby if the plot would damn their souls, to which Catesby had replied it would not, and that the plight of England's Catholics required that it be done. Catesby also apparently asked for £2,000, and the use of Rushton Hall in Northamptonshire. Tresham declined both offers (although he did give £100 to Thomas Wintour), and told his interrogators that he had moved his family from Rushton to London in advance of the plot; hardly the actions of a guilty man, he claimed.


title: Gunpowder Plot

text: The details of the plot were finalised in October, in a series of taverns across London and Daventry. Fawkes would be left to light the fuse and then escape across the Thames, while simultaneously a revolt in the Midlands would help to ensure the capture of the King's daughter, Elizabeth. Fawkes would leave for the continent, to explain events in England to the European Catholic powers.


title: Gunpowder Plot

text: The wives of those involved and Anne Vaux (a friend of Garnet who often shielded priests at her home) became increasingly concerned by what they suspected was about to happen. Several of the conspirators expressed worries about the safety of fellow Catholics who would be present in Parliament on the day of the planned explosion. Percy was concerned for his patron, Northumberland, and the young Earl of Arundel's name was brought up; Catesby suggested that a minor wound might keep him from the chamber on that day. The Lords Vaux, Montagu, Monteagle, and Stourton were also mentioned. Keyes suggested warning Lord Mordaunt, his wife's employer, to derision from Catesby.


title: Gunpowder Plot

text: On Saturday 26 October, Monteagle (Tresham's brother-in-law) arranged a meal in a long-disused house at Hoxton. Suddenly a servant appeared saying he had been handed a letter for Lord Monteagle from a stranger in the road. Monteagle ordered it to be read aloud to the company.


title: Gunpowder Plot

text: Uncertain of the letter's meaning, Monteagle promptly rode to Whitehall and handed it to Cecil (then Earl of Salisbury). Salisbury informed the Earl of Worcester, considered to have recusant sympathies, and the suspected Catholic Henry Howard, 1st Earl of Northampton, but kept news of the plot from the King, who was busy hunting in Cambridgeshire and not expected back for several days. Monteagle's servant, Thomas Ward, had family connections with the Wright brothers, and sent a message to Catesby about the betrayal. Catesby, who had been due to go hunting with the King, suspected that Tresham was responsible for the letter, and with Thomas Wintour confronted the recently recruited conspirator. Tresham managed to convince the pair that he had not written the letter, but urged them to abandon the plot. Salisbury was already aware of certain stirrings before he received the letter, but did not yet know the exact nature of the plot, or who exactly was involved. He therefore elected to wait, to see how events unfolded.


title: Gunpowder Plot

text: The letter was shown to the King on Friday 1 November following his arrival back in London. Upon reading it, James immediately seized upon the word "blow" and felt that it hinted at "some strategem of fire and powder", perhaps an explosion exceeding in violence the one that killed his father, Lord Darnley, at Kirk o' Field in 1567. Keen not to seem too intriguing, and wanting to allow the King to take the credit for unveiling the conspiracy, Salisbury feigned ignorance. The following day members of the Privy Council visited the King at the Palace of Whitehall and informed him that, based on the information that Salisbury had given them a week earlier, on Monday the Lord Chamberlain Thomas Howard, 1st Earl of Suffolk would undertake a search of the Houses of Parliament, "both above and below". On Sunday 3 November, Percy, Catesby and Wintour had a final meeting, where Percy told his colleagues that they should "abide the uttermost triall", and reminded them of their ship waiting at anchor on the Thames.


title: Gunpowder Plot

text: By 4 November, Digby was ensconced with a "hunting party" at Dunchurch, ready to abduct Elizabeth. The same day, Percy visited the Earl of Northumberland—who was uninvolved in the conspiracy—to see if he could discern what rumours surrounded the letter to Monteagle. Percy returned to London and assured Wintour, John Wright, and Robert Keyes that they had nothing to be concerned about, and returned to his lodgings on Gray's Inn Road. That same evening Catesby, likely accompanied by John Wright and Bates, set off for the Midlands. Fawkes visited Keyes, and was given a pocket watch left by Percy, to time the fuse, and an hour later Rookwood received several engraved swords from a local cutler.


title: Gunpowder Plot

text: Although two accounts of the number of searches and their timing exist, according to the King's version, the first search of the buildings in and around Parliament was made on Monday 4 November—as the plotters were busy making their final preparations—by Suffolk, Monteagle, and John Whynniard. They found a large pile of firewood in the undercroft beneath the House of Lords, accompanied by what they presumed to be a serving man (Fawkes), who told them that the firewood belonged to his master, Thomas Percy. They left to report their findings, at which time Fawkes also left the building. The mention of Percy's name aroused further suspicion as he was already known to the authorities as a Catholic agitator. The King insisted that a more thorough search be undertaken. Late that night, the search party, headed by Thomas Knyvet, returned to the undercroft. They again found Fawkes, dressed in a cloak and hat, and wearing boots and spurs. He was arrested, whereupon he gave his name as John Johnson. He was carrying a lantern now held in the Ashmolean Museum, Oxford, and a search of his person revealed a pocket watch, several slow matches and touchwood. 36 barrels of gunpowder were discovered hidden under piles of faggots and coal. Fawkes was taken to the King early on the morning of 5 November.


title: Gunpowder Plot

text: As news of "John Johnson's" arrest spread among the plotters still in London, most fled northwest, along Watling Street. Christopher Wright and Thomas Percy left together. Rookwood left soon after, and managed to cover 30 miles in two hours on one horse. He overtook Keyes, who had set off earlier, then Wright and Percy at Little Brickhill, before catching Catesby, John Wright, and Bates on the same road. Reunited, the group continued northwest to Dunchurch, using horses provided by Digby. Keyes went to Mordaunt's house at Drayton. Meanwhile, Thomas Wintour stayed in London, and even went to Westminster to see what was happening. When he realised the plot had been uncovered, he took his horse and made for his sister's house at Norbrook, before continuing to Huddington Court.


title: Gunpowder Plot

text: The group of six conspirators stopped at Ashby St Ledgers at about 6 pm, where they met Robert Wintour and updated him on their situation. They then continued on to Dunchurch, and met with Digby. Catesby convinced him that despite the plot's failure, an armed struggle was still a real possibility. He announced to Digby's "hunting party" that the King and Salisbury were dead, before the fugitives moved west to Warwick.


title: Gunpowder Plot

text: In London, news of the plot was spreading, and the authorities set extra guards on the city gates, closed the ports, and protected the house of the Spanish Ambassador, which was surrounded by an angry mob. An arrest warrant was issued against Thomas Percy, and his patron, the Earl of Northumberland, was placed under house arrest. In "John Johnson's" initial interrogation he revealed nothing other than the name of his mother, and that he was from Yorkshire. A letter to Guy Fawkes was discovered on his person, but he claimed that name was one of his aliases. Far from denying his intentions, "Johnson" stated that it had been his purpose to destroy the King and Parliament. Nevertheless, he maintained his composure and insisted that he had acted alone. His unwillingness to yield so impressed the King that he described him as possessing "a Roman resolution".


title: Gunpowder Plot

text: On 6 November, the Lord Chief Justice, Sir John Popham (a man with a deep-seated hatred of Catholics) questioned Rookwood's servants. By the evening he had learned the names of several of those involved in the conspiracy: Catesby, Rookwood, Keyes, Wynter, John and Christopher Wright, and Grant. "Johnson" meanwhile persisted with his story, and along with the gunpowder he was found with, was moved to the Tower of London, where the King had decided that "Johnson" would be tortured. The use of torture was forbidden, except by royal prerogative or a body such as the Privy Council or Star Chamber. In a letter of 6 November James wrote: "The gentler tortours [tortures] are to be first used unto him, [and thus by steps extended to the bottom depths], and so God speed your good work." "Johnson" may have been placed in manacles and hung from the wall, but he was almost certainly subjected to the horrors of the rack. On 7 November his resolve was broken; he confessed late that day, and again over the following two days.


title: Gunpowder Plot

text: On 6 November, with Fawkes maintaining his silence, the fugitives raided Warwick Castle for supplies, then continued to Norbrook to collect weapons. From there they continued their journey to Huddington. Bates left the group and travelled to Coughton Court to deliver a letter from Catesby, to Father Garnet and the other priests, informing them of what had transpired, and asking for their help in raising an army. Garnet replied by begging Catesby and his followers to stop their "wicked actions", before himself fleeing. Several priests set out for Warwick, worried about the fate of their colleagues. They were caught, and then imprisoned in London. Catesby and the others arrived at Huddington early in the afternoon, and were met by Thomas Wintour. They received practically no support or sympathy from those they met, including family members, who were terrified at the prospect of being associated with treason. They continued on to Holbeche House on the border of Staffordshire, the home of Stephen Littleton, a member of their ever-decreasing band of followers. Whilst there, Stephen Littleton and Thomas Wintour went to Pepperhill, the Shropshire residence at Boningale of Robert Wintour's father-in-law John Talbot, to gain support, but to no avail. Tired and desperate, they spread out some of the now-soaked gunpowder in front of the fire, to dry out. Although gunpowder does not explode unless physically contained, a spark from the fire landed on the powder and the resultant flames engulfed Catesby, Rookwood, Grant, and a man named Morgan, who was a member of the hunting party.


title: Gunpowder Plot

text: Thomas Wintour and Littleton, on their way from Huddington to Holbeche House, were told by a messenger that Catesby had died. At that point, Littleton left, but Thomas arrived at the house to find Catesby alive, albeit scorched. John Grant was not so lucky, and had been blinded by the fire. Digby, Robert Wintour and his half-brother John, and Thomas Bates, had all left. Of the plotters, only the singed figures of Catesby and Grant, the Wright brothers, Rookwood, and Percy remained. The fugitives resolved to stay in the house and wait for the arrival of the King's men.


title: Gunpowder Plot

text: Richard Walsh (Sheriff of Worcestershire) and his company of 200 men besieged Holbeche House on the morning of 8 November. Thomas Wintour was hit in the shoulder while crossing the courtyard. John Wright was shot, followed by his brother, and then Rookwood. Catesby and Percy were reportedly killed by a single lucky shot. The attackers rushed the property, and stripped the dead or dying defenders of their clothing. Grant, Morgan, Rookwood, and Wintour were arrested.


title: Gunpowder Plot

text: Bates and Keyes were captured shortly after Holbeche House was taken. Digby, who had intended to give himself up, was caught by a small group of pursuers. Tresham was arrested on 12 November, and taken to the Tower three days later. Montague, Mordaunt, and Stourton (Tresham's brother-in-law) were also imprisoned in the Tower. The Earl of Northumberland joined them on 27 November. Meanwhile, the government used the revelation of the plot to accelerate its persecution of Catholics. The home of Anne Vaux at Enfield Chase was searched, revealing the presence of trap doors and hidden passages. A terrified servant then revealed that Garnet, who had often stayed at the house, had recently given a Mass there. Father John Gerard was secreted at the home of Elizabeth Vaux, in Harrowden. Vaux was taken to London for interrogation. There she was resolute; she had never been aware that Gerard was a priest, she had presumed he was a "Catholic gentleman", and she did not know of his whereabouts. The homes of the conspirators were searched, and looted; Mary Digby's household was ransacked, and she was made destitute. Some time before the end of November, Garnet moved to Hindlip Hall near Worcester, the home of the Habingtons, where he wrote a letter to the Privy Council protesting his innocence.


title: Gunpowder Plot

text: The foiling of the Gunpowder Plot initiated a wave of national relief at the delivery of the King and his sons, and inspired in the ensuing parliament a mood of loyalty and goodwill, which Salisbury astutely exploited to extract higher subsidies for the King than any (bar one) granted in Elizabeth I's reign. Walter Raleigh, who was languishing in the Tower owing to his involvement in the Main Plot, and whose wife was a first cousin of Lady Catesby, declared he had had no knowledge of the conspiracy. The Bishop of Rochester gave a sermon at St. Paul's Cross, in which he condemned the plot. In his speech to both Houses on 9 November, James expounded on two emerging preoccupations of his monarchy: the divine right of kings and the Catholic question. He insisted that the plot had been the work of only a few Catholics, not of the English Catholics as a whole, and he reminded the assembly to rejoice at his survival, since kings were divinely appointed and he owed his escape to a miracle. Salisbury wrote to his English ambassadors abroad, informing them of what had occurred, and also reminding them that the King bore no ill will to his Catholic neighbours. The foreign powers largely distanced themselves from the plotters, calling them atheists and Protestant heretics.


title: Gunpowder Plot

text: Sir Edward Coke was in charge of the interrogations. Over a period of about ten weeks, in the Lieutenant's Lodgings at the Tower of London (now known as the Queen's House) he questioned those who had been implicated in the plot. For the first round of interrogations, no real proof exists that these people were tortured, although on several occasions Salisbury certainly suggested that they should be. Coke later revealed that the threat of torture was in most cases enough to elicit a confession from those caught up in the aftermath of the plot.


title: Gunpowder Plot

text: Only two confessions were printed in full: Fawkes's confession of 8 November, and Wintour's of 23 November. Having been involved in the conspiracy from the start (unlike Fawkes), Wintour was able to give extremely valuable information to the Privy Council. The handwriting on his testimony is almost certainly that of the man himself, but his signature was markedly different. Wintour had previously only ever signed his name as such, but his confession is signed "Winter", and since he had been shot in the shoulder, the steady hand used to write the signature may indicate some measure of government interference—or it may indicate that writing a shorter version of his name was less painful. Wintour's testimony makes no mention of his brother, Robert. Both were published in the so-called "King's Book", a hastily written official account of the conspiracy published in late November 1605.


title: Gunpowder Plot

text: Henry Percy, Earl of Northumberland, was in a difficult position. His midday dinner with Thomas Percy on 4 November was damning evidence against him, and after Thomas Percy's death there was nobody who could either implicate him or clear him. The Privy Council suspected that Northumberland would have been Princess Elizabeth's protector had the plot succeeded, but there was insufficient evidence to convict him. Northumberland remained in the Tower and on 27 June 1606 was finally charged with contempt. He was stripped of all public offices, fined £30,000 (about £ in 2022), and kept in the Tower until June 1621. The Lords Mordaunt and Stourton were tried in the Star Chamber. They were condemned to imprisonment in the Tower, where they remained until 1608, when they were transferred to the Fleet Prison. Both were also given significant fines.


title: Gunpowder Plot

text: Several other people not involved in the conspiracy, but known or related to the conspirators, were also questioned. Northumberland's brothers, Sir Allen and Sir Josceline Percy, were arrested. Lord Montagu had employed Fawkes at an early age, and had also met Catesby on 29 October, and was therefore of interest; he was released several months later. Agnes Wenman was from a Catholic family, and related to Elizabeth Vaux. She was examined twice but the charges against her were eventually dropped. Percy's secretary and later the controller of Northumberland's household, Dudley Carleton, had leased the vault where the gunpowder was stored, and consequently he was imprisoned in the Tower. Salisbury believed his story, and authorised his release.


title: Gunpowder Plot

text: Thomas Bates confessed on 4 December, providing much of the information that Salisbury needed to link the Catholic clergy to the plot. Bates had been present at most of the conspirators' meetings, and under interrogation he implicated Father Tesimond in the plot. On 13 January 1606, he described how he had visited Garnet and Tesimond on 7 November to inform Garnet of the plot's failure. Bates also told his interrogators of his ride with Tesimond to Huddington, before the priest left him to head for the Habingtons at Hindlip Hall, and of a meeting between Garnet, Gerard, and Tesimond in October 1605.


title: Gunpowder Plot

text: At about the same time in December, Tresham's health began to deteriorate. He was visited regularly by his wife, a nurse, and his servant William Vavasour—who documented his strangury. Before he died, Tresham had also told of Garnet's involvement with the 1603 mission to Spain, but in his last hours he retracted some of these statements. Nowhere in his confession did he mention the Monteagle letter. He died early on the morning of 23 December, and was buried in the Tower. Nevertheless, he was attainted along with the other plotters; his head was set on a pike either (accounts differ) at Northampton or London Bridge, and his estates confiscated.


title: Gunpowder Plot

text: On 15 January a proclamation named Father Garnet, Father Gerard, and Father Greenway (Tesimond) as wanted men. Tesimond and Gerard escaped the country and lived out their days in freedom. Several days earlier, on 9 January, Robert Wintour and Stephen Littleton were captured. They had been hiding at Hagley, the home of Humphrey Littleton, brother of MP John Littleton, imprisoned for treason in 1601 for his part in the Essex revolt. They were betrayed by a cook, who grew suspicious of the amount of food sent up for his master's consumption. Humphrey denied the presence of the two fugitives, but another servant led the authorities to their hiding place. On 20 January, the local Justice of the Peace and his retainers arrived at Thomas Habington's home, Hindlip Hall, to arrest the Jesuits. Despite Thomas Habington's protests, the men spent the next four days searching the house. On 24 January, starving, two priests left their hiding places and were discovered. Humphrey Littleton, who had escaped from the authorities at Hagley, got as far as Prestwood in Staffordshire before he was captured. He was imprisoned, and then condemned to death at Worcester. On 26 January, in exchange for his life, he told the authorities where they could find Father Garnet. Worn down by hiding for so long, Garnet, accompanied by another priest, emerged from his priest hole the next day.


title: Gunpowder Plot

text: By coincidence, on the same day that Garnet was found, the surviving conspirators were arraigned in Westminster Hall. Seven of the prisoners were taken from the Tower to the Star Chamber by barge. Bates, who was considered lower class, was brought from the Gatehouse Prison. Some of the prisoners were reportedly despondent, but others were nonchalant, even smoking tobacco. The King and his family, hidden from view, were among the many who watched the trial. The Lords Commissioners present were the Earls of Suffolk, Worcester, Northampton, Devonshire, and Salisbury. Sir John Popham was Lord Chief Justice, Sir Thomas Fleming was Lord Chief Baron of the Exchequer, and two Justices, Sir Thomas Walmsley and Sir Peter Warburton, sat as Justices of the Common Pleas. The list of traitors' names was read aloud, beginning with those of the priests: Garnet, Tesimond, and Gerard.


title: Gunpowder Plot

text: The first to speak was the Speaker of the House of Commons (later Master of the Rolls), Sir Edward Philips, who described the intent behind the plot in lurid detail. He was followed by the Attorney-General Sir Edward Coke, who began with a long speech—the content of which was heavily influenced by Salisbury—that included a denial that the King had ever made any promises to the Catholics. Monteagle's part in the discovery of the plot was welcomed, and denunciations of the 1603 mission to Spain featured strongly. Fawkes's protestations that Gerard knew nothing of the plot were omitted from Coke's speech. The foreign powers, when mentioned, were accorded due respect, but the priests were accursed, their behaviour analysed and criticised wherever possible. There was little doubt, according to Coke, that the plot had been invented by the Jesuits. Garnet's meeting with Catesby, at which the former was said to have absolved the latter of any blame in the plot, was proof enough that the Jesuits were central to the conspiracy; according to Coke the Gunpowder Plot would always be known as the Jesuit Treason. Coke spoke with feeling of the probable fate of the Queen and the rest of the King's family, and of the innocents who would have been caught up in the explosion.


title: Gunpowder Plot

text: Each of the condemned, said Coke, would be drawn backwards to his death, by a horse, his head near the ground. He was to be "put to death halfway between heaven and earth as unworthy of both". His genitals would be cut off and burnt before his eyes, and his bowels and heart then removed. Then he would be decapitated, and the dismembered parts of his body displayed so that they might become "prey for the fowls of the air". Confessions and declarations from the prisoners were then read aloud, and finally the prisoners were allowed to speak. Rookwood claimed that he had been drawn into the plot by Catesby, "whom he loved above any worldy man". Thomas Wintour begged to be hanged for himself and his brother, so that his brother might be spared. Fawkes explained his not guilty plea as ignorance of certain aspects of the indictment. Keyes appeared to accept his fate, Bates and Robert Wintour begged for mercy, and Grant explained his involvement as "a conspiracy intended but never effected". Only Digby, tried on a separate indictment, pleaded guilty, insisting that the King had reneged upon promises of toleration for Catholics, and that affection for Catesby and love of the Catholic cause mitigated his actions. He sought death by the axe and begged mercy from the King for his young family. His defence was in vain; his arguments were rebuked by Coke and Northumberland, and along with his seven co-conspirators, he was found guilty by the jury of high treason. Digby shouted "If I may but hear any of your lordships say, you forgive me, I shall go more cheerfully to the gallows." The response was short: "God forgive you, and we do."


title: Gunpowder Plot

text: Garnet may have been questioned on as many as 23 occasions. His response to the threat of the rack was " [Threats are only for boys]", and he denied having encouraged Catholics to pray for the success of the "Catholic Cause". His interrogators resorted to the forgery of correspondence between Garnet and other Catholics, but to no avail. His jailers then allowed him to talk with another priest in a neighbouring cell, with eavesdroppers listening to every word. Eventually Garnet let slip a crucial piece of information, that there was only one man who could testify that he had any knowledge of the plot. Under torture Garnet admitted that he had heard of the plot from fellow Jesuit Oswald Tesimond, who had learnt of it in confession from Catesby. Garnet was charged with high treason and tried in the Guildhall on 28 March, in a trial lasting from 8 am until 7 pm. According to Coke, Garnet instigated the plot: "[Garnet] hath many gifts and endowments of nature, by art learned, a good linguist and, by profession, a Jesuit and a Superior as indeed he is Superior to all his predecessors in devilish treason, a Doctor of Dissimulation, Deposing of Princes, Disposing of Kingdoms, Daunting and deterring of subjects, and Destruction." Garnet refuted all the charges against him, and explained the Catholic position on such matters, but he was nevertheless found guilty and sentenced to death.


title: Gunpowder Plot

text: Although Catesby and Percy escaped the executioner, their bodies were exhumed and decapitated, and their heads exhibited on spikes outside the House of Lords. On a cold 30 January, Everard Digby, Robert Wintour, John Grant, and Thomas Bates were tied to hurdles—wooden panels—and dragged through the crowded streets of London to St Paul's Churchyard. Digby, the first to mount the scaffold, asked the spectators for forgiveness, and refused the attentions of a Protestant clergyman. He was stripped of his clothing, and wearing only a shirt, climbed the ladder to place his head through the noose. He was quickly cut down, and while still fully conscious was castrated, disembowelled, and then quartered, along with the three other prisoners. The following day, Thomas Wintour, Ambrose Rookwood, Robert Keyes, and Guy Fawkes were hanged, drawn and quartered, opposite the building they had planned to blow up, in the Old Palace Yard at Westminster. Keyes did not wait for the hangman's command and jumped from the gallows, but he survived the drop and was led to the quartering block. Although weakened by his torture, Fawkes managed to jump from the gallows and break his neck, thus avoiding the agony of the gruesome latter part of his execution.


title: Gunpowder Plot

text: Steven Littleton was executed at Stafford. His cousin Humphrey, despite his co-operation with the authorities, met his end at Red Hill near Worcester. Henry Garnet's execution took place on 3 May 1606.


title: Gunpowder Plot

text: Greater freedom for Roman Catholics to worship as they chose seemed unlikely in 1604, but the discovery of such a wide-ranging conspiracy, the capture of those involved, and the subsequent trials, led Parliament to consider introducing new anti-Catholic legislation. The event also destroyed all hope that the Spanish would ever secure tolerance of the Catholics in England. In the summer of 1606, laws against recusancy were strengthened; the Popish Recusants Act returned England to the Elizabethan system of fines and restrictions, introduced a sacramental test, and an Oath of Allegiance, requiring Catholics to abjure as a "heresy" the doctrine that "princes excommunicated by the Pope could be deposed or assassinated". Catholic emancipation took another 200 years, but many important and loyal Catholics retained high office during King James I's reign. Although there was no "golden time" of "toleration" of Catholics, which Father Garnet had hoped for, James's reign was nevertheless a period of relative leniency for Catholics, and few were subject to prosecution.


title: Gunpowder Plot

text: The playwright William Shakespeare had already used the family history of Northumberland's family in his "Henry IV" series of plays, and the events of the Gunpowder Plot seem to have featured alongside the earlier Gowrie conspiracy in "Macbeth", written some time between 1603 and 1607. Interest in the demonic was heightened by the Gunpowder Plot. The King had become engaged in the great debate about other-worldly powers in writing his "Daemonologie" in 1599, before he became King of England as well as Scotland. Inversions seen in such lines as "fair is foul and foul is fair" are used frequently, and another possible reference to the plot relates to the use of equivocation; Garnet's "A Treatise of Equivocation" was found on one of the plotters. Another writer influenced by the plot was John Milton, who in 1626 wrote what one commentator has called a "critically vexing poem", "In Quintum Novembris". Reflecting "partisan public sentiment on an English-Protestant national holiday", in the published editions of 1645 and 1673, the poem is preceded by five epigrams on the subject of the Gunpowder Plot, apparently written by Milton in preparation for the larger work. The plot may also have influenced his later work, "Paradise Lost".


title: Gunpowder Plot

text: The Gunpowder Plot was commemorated for years by special sermons and other public acts, such as the ringing of church bells. It added to an increasingly full calendar of Protestant celebrations that contributed to the national and religious life of 17th-century England, and has evolved into the Bonfire Night of today. In "What If the Gunpowder Plot Had Succeeded?" historian Ronald Hutton considered the events which might have followed a successful implementation of the plot; the destruction of the House of Lords and all those within it. He concluded that a severe backlash against suspected Catholics would have followed, and that without foreign assistance a successful rebellion would have been unlikely; despite differing religious convictions, most Englishmen were loyal to the institution of the monarchy. England might have become a more "Puritan absolute monarchy", as "existed in Sweden, Denmark, Saxony, and Prussia in the seventeenth century", rather than following the path of parliamentary and civil reform that it did.


title: Gunpowder Plot

text: Many at the time felt that Salisbury had been involved in the plot to gain favour with the King and enact more stridently anti-Catholic legislation. Such conspiracy theories alleged that Salisbury had either actually invented the plot or allowed it to continue when his agents had already infiltrated it, for the purposes of propaganda. The Popish Plot of 1678 sparked renewed interest in the Gunpowder Plot, resulting in a book by Thomas Barlow, Bishop of Lincoln, which refuted "a bold and groundless surmise that all this was a contrivance of Secretary Cecil".


title: Gunpowder Plot

text: In 1897 Father John Gerard of Stonyhurst College, namesake of John Gerard (who, following the plot's discovery, had evaded capture), wrote an account called "What was the Gunpowder Plot?", alleging Salisbury's culpability. This prompted a refutation later that year by Samuel Gardiner, who argued that Gerard had gone too far in trying to "wipe away the reproach" which the plot had exacted on generations of English Catholics. Gardiner portrayed Salisbury as guilty of nothing more than opportunism. Subsequent attempts to prove Salisbury's involvement, such as Francis Edwards's 1969 work "Guy Fawkes: the real story of the gunpowder plot?", have similarly foundered on the lack of any clear evidence.


title: Gunpowder Plot

text: The cellars under the Houses of Parliament continued to be leased out to private individuals until 1678, when news of the Popish Plot broke. It was then considered prudent to search the cellars on the day before each State Opening of Parliament, a ritual that survives to this day.


title: Gunpowder Plot

text: In January 1606, during the first sitting of Parliament since the plot, the Observance of 5th November Act 1605 was passed, making services and sermons commemorating the event an annual feature of English life; the act remained in force until 1859. The tradition of marking the day with the ringing of church bells and bonfires started soon after the Plot's discovery, and fireworks were included in some of the earliest celebrations. In Britain, 5 November is variously called Bonfire Night, Fireworks Night, or Guy Fawkes Night.


title: Gunpowder Plot

text: 5 November firework displays and bonfire parties are common throughout Britain, both in major public displays and in private gardens. Traditionally, in the weeks running up to the 5th, children made "guys"—effigies supposedly of Fawkes—usually made from old clothes stuffed with newspaper, and fitted with a grotesque mask, to be burnt on 5 November bonfire. These "guys" were exhibited in the street to collect money for fireworks, although this custom has become less common. The word "guy", in the 19th century, thus came to mean an oddly dressed person and, in the 20th and 21st centuries, to mean any male person.


title: Gunpowder Plot

text: According to the biographer Esther Forbes, the Guy Fawkes Day celebration in the pre-revolutionary American colonies was a very popular holiday. In Boston, the revelry on "Pope Night" took on anti-authoritarian overtones, and often became so dangerous that many would not venture out of their homes.


title: Gunpowder Plot

text: In the 2005, ITV programme "", a full-size replica of the House of Lords was built and destroyed with barrels of gunpowder, totalling 1 tonne of explosives. The experiment was conducted on the Advantica-owned Spadeadam test site and demonstrated that the explosion—if the gunpowder had been in good order—would have killed all those in the building. The power of the explosion was such that of the deep concrete walls making up the undercroft (replicating how archives suggest the walls of the old House of Lords were constructed), the end wall where the barrels were placed, under the throne, was reduced to rubble, and the adjacent surviving portions of wall were shoved away. Measuring devices placed in the chamber to calculate the force of the blast were recorded as going off the scale just before their destruction by the explosion; a piece of the head of the dummy representing King James, which had been placed on a throne inside the chamber—surrounded by courtiers, peers and bishops—was found a considerable distance from its initial location. According to the findings of the programme, no one within of the blast could have survived, and all of the stained glass windows in Westminster Abbey would have shattered, as would all of the windows in the vicinity of the Palace. The explosion would have been seen from miles away and heard from further away still. Even if only half of the gunpowder had gone off—which Fawkes was apparently prepared for—everyone in the House of Lords and its environs would have been killed instantly.


title: Gunpowder Plot

text: The programme also disproved claims that some deterioration in the quality of the gunpowder would have prevented the explosion. A portion of deliberately deteriorated gunpowder, of such low quality as to make it unusable in firearms, when placed in a heap and ignited, still managed to create a large explosion. The impact of even deteriorated gunpowder would have been magnified by its containment in wooden barrels, compensating for the quality of the contents. The compression would have created a cannon effect, with the powder first blowing up from the top of the barrel before, a millisecond later, blowing out. Calculations showed that Fawkes, who was skilled in the use of gunpowder, had deployed double the amount needed. In


title: Gunpowder Plot

text: n a test detonation of all of period-accurate gunpowder available in the UK inside the same size of barrel Fawkes had used, the experts for the project were surprised at how much more powerful an effect compression had in creating an explosion.


title: Gunpowder Plot

text: Some of the gunpowder guarded by Fawkes may have survived. In March 2002, workers cataloguing archives of diarist John Evelyn at the British Library found a box containing a number of gunpowder samples, including a compressed bar with a note in Evelyn's handwriting stating that it had belonged to Guy Fawkes. A further note, written in the 19th century, confirmed this provenance, although in 1952 the document acquired a new comment: "but there was none left!"


title: State religion

text: A state religion (also called religious state or official religion) is a religion or creed officially endorsed by a sovereign state. A state with an official religion (also known as confessional state), while not secular, is not necessarily a theocracy. State religions are official or government-sanctioned establishments of a religion, but the state does not need to be under the control of the religion (as in a theocracy) nor is the state-sanctioned religion necessarily under the control of the state.


title: State religion

text: Official religions have been known throughout human history in almost all types of cultures, reaching into the Ancient Near East and prehistory. The relation of religious cult and the state was discussed by the ancient Latin scholar Marcus Terentius Varro, under the term of "theologia civilis" (). The first state-sponsored Christian church was the Armenian Apostolic Church, established in 301 CE. In Christianity, as the term "church" is typically applied to a place of worship for Christians or organizations incorporating such ones, the term "state church" is associated with Christianity as sanctioned by the government, historically the state church of the Roman Empire in the last centuries of the Empire's existence, and is sometimes used to denote a specific modern national branch of Christianity. Closely related to state churches are ecclesiae, which are similar but carry a more minor connotation.


title: State religion

text: In the Middle East, the majority of states with a predominantly Muslim population have Islam as their official religion, though the degree of religious restrictions on citizens' everyday lives varies by country. Rulers of Saudi Arabia use both secular and religious power, while Iran's secular presidents are supposed to follow the decisions of religious authorities since the 1979 Islamic Revolution. Turkey, which also has Muslim-majority population, became a secular country after Atatürk's Reforms, although unlike the Russian Revolution of the same time period, it did not result in the adoption of state atheism.


title: State religion

text: The degree to which an official national religion is imposed upon citizens by the state in contemporary society varies considerably; from high as in Saudi Arabia and Iran to minimal or none at all as Denmark, England, Iceland, and Greece.