document_title: Serotonin

question_text: where does serotonin come from in the brain

text: Serotonin

text: Functions[edit]

text: Serotonin is a neurotransmitter and is found in all bilateral animals including insects.[18] Serotonin is also present in plants (phytoserotonin).[19]

text: Perception of resource availability[edit]

text: Serotonin mediates the animal's perceptions of resource; In less complex animals, such as some invertebrates, resources simply mean food availability.[20] In plants serotonin synthesis seems to be associated with stress signals.[19] In more complex animals, such as arthropods and vertebrates, resources also can mean social dominance.[21] In response to the perceived abundance or scarcity of resources, an animal's growth, reproduction or mood may be elevated or lowered. This may somewhat depend on how much serotonin the organism has at its disposal.[citation needed]

text: Cellular effects[edit]

text: Receptors[edit]

text: The 5-HT receptors, the receptors for serotonin, are located on the cell membrane of nerve cells and other cell types in animals, and mediate the effects of serotonin as the endogenous ligand and of a broad range of pharmaceutical and hallucinogenic drugs. Except for the 5-HT3 receptor, a ligand-gated ion channel, all other 5-HT receptors are G-protein-coupled receptors (also called seven-transmembrane, or heptahelical receptors) that activate an intracellular second messenger cascade.[22]

text: Termination[edit]

text: Serotonergic action is terminated primarily via uptake of 5-HT from the synapse. This is accomplished through the specific monoamine transporter for 5-HT, SERT, on the presynaptic neuron. Various agents can inhibit 5-HT reuptake, including cocaine, dextromethorphan (an antitussive), tricyclic antidepressants and selective serotonin reuptake inhibitors (SSRIs). A 2006 study conducted by the University of Washington suggested that a newly discovered monoamine transporter, known as PMAT, may account for "a significant percentage of 5-HT clearance".[23]

text: Contrasting with the high-affinity SERT, the PMAT has been identified as a low-affinity transporter, with an apparent Km of 114 micromoles/l for serotonin; approximately 230 times higher than that of SERT. However, the PMAT, despite its relatively low serotonergic affinity, has a considerably higher transport 'capacity' than SERT, "resulting in roughly comparable uptake efficiencies to SERT in heterologous expression systems."[this quote needs a citation] The study also suggests some SSRIs, such as fluoxetine and sertraline anti-depressants, inhibit PMAT but at IC50 values which surpass the therapeutic plasma concentrations by up to four orders of magnitude. Therefore, SSRI monotherapy is "ineffective" in PMAT inhibition. At present, no known pharmaceuticals are known to appreciably inhibit PMAT at normal therapeutic doses. The PMAT also suggestively transports dopamine and norepinephrine, albeit at Km values even higher than that of 5-HT (330–15,000 μmoles/L).[citation needed]

text: Serotonylation[edit]

text: Serotonin can also signal through a nonreceptor mechanism called serotonylation, in which serotonin modifies proteins.[24] This process underlies serotonin's effects upon platelet-forming cells (thrombocytes) in which it links to the modification of signaling enzymes called GTPases that then trigger the release of vesicle contents by exocytosis.[25] A similar process underlies the pancreatic release of insulin.[24]

text: The effects of serotonin upon vascular smooth muscle tone (this is the biological function from which serotonin originally got its name) depend upon the serotonylation of proteins involved in the contractile apparatus of muscle cells.[26]

text:
Binding profile of serotonin

Receptor
Ki (nM)[27]
Receptor function[Note 1]

5-HT1 receptor family signals via Gi/o inhibition of adenylyl cyclase.

5-HT1A
3.17
Memory[vague] (agonists ↓); learning[vague] (agonists ↓); anxiety (agonists ↓); depression (agonists ↓); positive, negative, and cognitive symptoms of schizophrenia (partial agonists ↓); analgesia (agonists ↑); aggression (agonists ↓); dopamine release in the prefrontal cortex (agonists ↑); serotonin release and synthesis (agonists ↓)

5-HT1B
4.32
Vasoconstriction (agonists ↑); aggression (agonists ↓); bone mass (↓). Serotonin autoreceptor.

5-HT1D
5.03
Vasoconstriction (agonists ↑)

5-HT1E
7.53

5-HT1F
10

5-HT2 receptor family signals via Gq activation of phospholipase C.

5-HT2A
11.55
Psychedelia (agonists ↑); depression (agonists & antagonists ↓); anxiety (antagonists ↓); positive and negative symptoms of schizophrenia (antagonists ↓); norepinephrine release from the locus coeruleus (antagonists ↑); glutamate release in the prefrontal cortex (agonists ↑); urinary bladder contractions (agonists ↑)[28]

5-HT2B
8.71
Cardiovascular functioning (agonists increase risk of pulmonary hypertension), empathy (via the spindle neurons or Von Economo neurons[29])

5-HT2C
5.02
Dopamine release into the mesocorticolimbic pathway (agonists ↓); acetylcholine release in the prefrontal cortex (agonists ↑); appetite (agonists ↓); antipsychotic effects (agonists ↑); antidepressant effects (agonists & antagonists ↑)

Other 5-HT receptors

5-HT3
593
Emesis (agonists ↑); anxiolysis (antagonists ↑).

5-HT4
125.89
Movement of food across the GI tract (agonists ↑); memory & learning (agonists ↑); antidepressant effects (agonists ↑). Signalling via Gαs activation of adenylyl cyclase.

5-HT5A
251.2
Memory consolidation.[30] Signals via Gi/o inhibition of adenylyl cyclase.

5-HT6
98.41
Cognition (antagonists ↑); antidepressant effects (agonists & antagonists ↑); anxiogenic effects (antagonists ↑[31]). Gs signalling via activating adenylyl cyclase.

5-HT7
8.11
Cognition (antagonists ↑); antidepressant effects (antagonists ↑). Acts by Gs signalling via activating adenylyl cyclase.

text: Nervous system[edit]

text: The neurons of the raphe nuclei are the principal source of 5-HT release in the brain.[32] There are nine raphe nuclei, designated B1-B9, which contain the majority of serotonin-containing neurons (some scientists chose to group the nuclei raphes lineares into one nucleus), all of which are located along the midline of the brainstem, and centered on the reticular formation.[33][34] Axons from the neurons of the raphe nuclei form a neurotransmitter system reaching almost every part of the central nervous system. Axons of neurons in the lower raphe nuclei terminate in the cerebellum and spinal cord, while the axons of the higher nuclei spread out in the entire brain.

text: Ultrastructure and Function[edit]

text: The serotonin nuclei may also be divided into two main groups, the rostral and caudal containing three and four nuclei respectively. The rostral group consists of the caudal linear nuclei (B8), the dorsal raphe nuclei (B6 and B7) and the median raphe nuclei (B5, B8 and B9), that project into multiple cortical and subcortical structures. The caudal group consists of the nucleus raphe magnus (B3), raphe obscurus nucleus (B2), raphe pallidus nucleus (B1), and lateral medullary reticular formation, that project into the brainstem.[35]

text: Serotonergic pathway are involved in sensorimotor function, with pathways projecting both into cortical (Dorsal and Median Raphe Nuclei), subcortical, and spinal areas involved in motor activity. Pharmacological manipulation suggest that serotonergic activity increases with motor activity, while firing rates of serotonergic neurons increase with intense visual stimuli. The descending projections form a pathway that inhibits pain called the "descending inhibitory pathway" that may be relevant to disorder such as fibromyalgia, migraine and other pain disorders, and the efficacy of antidepressants in them.[36]

text: Serotonergic projections from the caudal nuclei are involved in regulating mood, emotion and hypo[37] or hyperserotonergic[38] states may be involved in depression and sickness behavior.

text: Microanatomy[edit]

text: Serotonin is released into the synapse, or space between neurons, and diffuses over a relatively wide gap (>20 µm) to activate 5-HT receptors located on the dendrites, cell bodies and presynaptic terminals of adjacent neurons.

text: When humans smell food, dopamine is released to increase the appetite. But, unlike in worms, serotonin does not increase anticipatory behaviour in humans; instead, the serotonin released while consuming activates 5-HT2C receptors on dopamine-producing cells. This halts their dopamine release, and thereby serotonin decreases appetite. Drugs that block 5-HT2C receptors make the body unable to recognize when it is no longer hungry or otherwise in need of nutrients, and are associated with weight gain,[39] especially in people with a low number of receptors.[40] The expression of 5-HT2C receptors in the hippocampus follows a diurnal rhythm,[41] just as the serotonin release in the ventromedial nucleus, which is characterised by a peak at morning when the motivation to eat is strongest.[42]

text: The amount of food that an animal acquires depends not only on food availability but also on the animal's ability to compete with others. This is especially true for social animals, where the stronger individuals might steal food from the weaker (this is not to say some non-social animals do not concern themselves with the needs of others or steal food from others[citation needed]). Thus, serotonin is involved not only in the perception of food availability but also in social rank.

text: In macaques, alpha males have twice the level of serotonin released in the brain than subordinate males and females (as measured by the levels of 5-Hydroxyindoleacetic acid (5-HIAA) in the cerebro-spinal fluid). Dominance status and cerebro-serotonin levels appear to be positively correlated. When dominant males were removed from such groups, subordinate males begin competing for dominance. Once new dominance hierarchies were established, serotonin levels of the new dominant individuals also increased to double those in subordinate males and females. The reason why serotonin levels are only high in dominant males but not dominant females has not yet been established.[43]

text: In humans, levels of 5-HT1A receptor activation in the brain show negative correlation with aggression,[44] and a mutation in the gene that codes for the 5-HT2A receptor may double the risk of suicide for those with that genotype.[45] Serotonin in the brain is not usually degraded after use, but is collected by serotonergic neurons by serotonin transporters on their cell surfaces. Studies have revealed nearly 10% of total variance in anxiety-related personality depends on variations in the description of where, when and how many serotonin transporters the neurons should deploy.[46]

text: Psychological influences[edit]

text: Serotonin has been implicated in cognition, mood, anxiety and psychosis, but strong clarity has not been achieved.[47][48]

text: Outside the nervous system[edit]

text: In the digestive tract (emetic)[edit]

text: The gut is surrounded by enterochromaffin cells, which release serotonin in response to food in the lumen. This makes the gut contract around the food. Platelets in the veins draining the gut collect excess serotonin.

text: If irritants are present in the food, the enterochromaffin cells release more serotonin to make the gut move faster, i.e., to cause diarrhea, so the gut is emptied of the noxious substance. If serotonin is released in the blood faster than the platelets can absorb it, the level of free serotonin in the blood is increased. This activates 5-HT3 receptors in the chemoreceptor trigger zone that stimulate vomiting.[49] The enterochromaffin cells not only react to bad food but are also very sensitive to irradiation and cancer chemotherapy. Drugs that block 5HT3 are very effective in controlling the nausea and vomiting produced by cancer treatment, and are considered the gold standard for this purpose.[50]

text: Bone metabolism[edit]

text: In mice and humans, alterations in serotonin levels and signalling have been shown to regulate bone mass.[51][52][53][54] Mice that lack brain serotonin have osteopenia, while mice that lack gut serotonin have high bone density. In humans, increased blood serotonin levels have been shown to be significant negative predictor of low bone density. Serotonin can also be synthesized, albeit at very low levels, in the bone cells. It mediates its actions on bone cells using three different receptors. Through 5-HT1B receptors, it negatively regulates bone mass, while it does so positively through 5-HT2B receptors and 5-HT2C receptors. There is very delicate balance between physiological role of gut serotonin and its pathology. Increase in the extracellular content of serotonin results in a complex relay of signals in the osteoblasts culminating in FoxO1/ Creb and ATF4 dependent transcriptional events.[55] These studies have opened a new area of research in bone metabolism that can be potentially harnessed to treat bone mass disorders.[56]

text: Organ development[edit]

text: Since serotonin signals resource availability it is not surprising that it affects organ development. Many human and animal studies have shown that nutrition in early life can influence, in adulthood, such things as body fatness, blood lipids, blood pressure, atherosclerosis, behavior, learning and longevity.[57][58][59] Rodent experiment shows that neonatal exposure to SSRI's makes persistent changes in the serotonergic transmission of the brain resulting in behavioral changes,[60][61] which are reversed by treatment with antidepressants.[62] By treating normal and knockout mice lacking the serotonin transporter with fluoxetine scientists showed that normal emotional reactions in adulthood, like a short latency to escape foot shocks and inclination to explore new environments were dependent on active serotonin transporters during the neonatal period.[63][64]

text: Human serotonin can also act as a growth factor directly. Liver damage increases cellular expression of 5-HT2A and 5-HT2B receptors, mediating liver compensatory regrowth (see Liver § Regeneration and transplantation)[65] Serotonin present in the blood then stimulates cellular growth to repair liver damage.[66] 5HT2B receptors also activate osteocytes, which build up bone[67] However, serotonin also inhibits osteoblasts, through 5-HT1B receptors.[68]

text: Cardiovascular growth factor[edit]

text: Serotonin, in addition, evokes endothelial nitric oxide synthase activation and stimulates, through a 5-HT1B receptor-mediated mechanism, the phosphorylation of p44/p42 mitogen-activated protein kinase activation in bovine aortic endothelial cell cultures.[69] In blood, serotonin is collected from plasma by platelets, which store it. It is thus active wherever platelets bind in damaged tissue, as a vasoconstrictor to stop bleeding, and also as a fibrocyte mitotic (growth factor), to aid healing.[70]

text: Pharmacology[edit]

text: Several classes of drugs target the 5-HT system, including some antidepressants, antipsychotics, anxiolytics, antiemetics, and antimigraine drugs, as well as the psychedelic drugs and empathogens.

text: Psychedelic drugs[edit]

text: The psychedelic drugs psilocin/psilocybin, DMT, mescaline, psychedelic mushroom and LSD are agonists, primarily at 5HT2A/2C receptors.[71][72][73] The empathogen-entactogen MDMA releases serotonin from synaptic vesicles of neurons.[74]

text: Antidepressants[edit]

text: Drugs that alter serotonin levels are used in treating depression, generalized anxiety disorder and social phobia. Monoamine oxidase inhibitors (MAOIs) prevent the breakdown of monoamine neurotransmitters (including serotonin), and therefore increase concentrations of the neurotransmitter in the brain. MAOI therapy is associated with many adverse drug reactions, and patients are at risk of hypertensive emergency triggered by foods with high tyramine content, and certain drugs. Some drugs inhibit the re-uptake of serotonin, making it stay in the synaptic cleft longer. The tricyclic antidepressants (TCAs) inhibit the reuptake of both serotonin and norepinephrine. The newer selective serotonin reuptake inhibitors (SSRIs) have fewer side-effects and fewer interactions with other drugs.[75]

text: Certain SSRI medications have been shown to lower serotonin levels below the baseline after chronic use, despite initial increases.[76] The 5-HTTLPR gene codes for the number of serotonin transporters in the brain, with more serotonin transporters causing decreased duration and magnitude of serotonergic signaling.[77] The 5-HTTLPR polymorphism (l/l) causing more serotonin transporters to be formed is also found to be more resilient against depression and anxiety.[78][79]

text: Serotonin syndrome[edit]

text: Extremely high levels of serotonin can cause a condition known as serotonin syndrome, with toxic and potentially fatal effects. In practice, such toxic levels are essentially impossible to reach through an overdose of a single antidepressant drug, but require a combination of serotonergic agents, such as an SSRI with an MAOI.[80] The intensity of the symptoms of serotonin syndrome vary over a wide spectrum, and the milder forms are seen even at nontoxic levels.[81]

text: Antiemetics[edit]

text: Some 5-HT3 antagonists, such as ondansetron, granisetron, and tropisetron, are important antiemetic agents. They are particularly important in treating the nausea and vomiting that occur during anticancer chemotherapy using cytotoxic drugs. Another application is in the treatment of postoperative nausea and vomiting.

text: Other[edit]

text: Some serotonergic agonist drugs cause fibrosis anywhere in the body, particularly the syndrome of retroperitoneal fibrosis, as well as cardiac valve fibrosis.[82] In the past, three groups of serotonergic drugs have been epidemiologically linked with these syndromes. These are the serotonergic vasoconstrictive antimigraine drugs (ergotamine and methysergide),[82] the serotonergic appetite suppressant drugs (fenfluramine, chlorphentermine, and aminorex), and certain anti-Parkinsonian dopaminergic agonists, which also stimulate serotonergic 5-HT2B receptors. These include pergolide and cabergoline, but not the more dopamine-specific lisuride.[83]

text: As with fenfluramine, some of these drugs have been withdrawn from the market after groups taking them showed a statistical increase of one or more of the side effects described. An example is pergolide. The drug was declining in use since it was reported in 2003 to be associated with cardiac fibrosis.[84]

text: Two independent studies published in the New England Journal of Medicine in January 2007, implicated pergolide, along with cabergoline, in causing valvular heart disease.[85][86] As a result of this, the FDA removed pergolide from the United States market in March 2007.[87] (Since cabergoline is not approved in the United States for Parkinson's Disease, but for hyperprolactinemia, the drug remains on the market. Treatment for hyperprolactinemia requires lower doses than that for Parkinson's Disease, diminishing the risk of valvular heart disease).[88]

text: Methyl-tryptamines and hallucinogens[edit]

text: Several plants contain serotonin together with a family of related tryptamines that are methylated at the amino (NH2) and (OH) groups, are N-oxides, or miss the OH group. These compounds do reach the brain, although some portion of them are metabolized by monoamine oxidase enzymes (mainly MAO-A) in the liver. Examples are plants from the Anadenanthera genus that are used in the hallucinogenic yopo snuff. These compounds are widely present in the leaves of many plants, and may serve as deterrents for animal ingestion. Serotonin occurs in several mushrooms of the genus Panaeolus.[89]

text: Comparative biology and evolution[edit]

text: Unicellular organisms[edit]

text: Serotonin is used by a variety of single-cell organisms for various purposes. SSRIs have been found to be toxic to algae.[90] The gastrointestinal parasite Entamoeba histolytica secretes serotonin, causing a sustained secretory diarrhea in some people.[16][91] Patients infected with E. histolytica have been found to have highly elevated serum serotonin levels, which returned to normal following resolution of the infection.[92] E. histolytica also responds to the presence of serotonin by becoming more virulent.[93] This means serotonin secretion not only serves to increase the spread of enteamoebas by giving the host diarrhea but also serves to coordinate their behaviour according to their population density, a phenomenon known as quorum sensing. Outside the gut of a host, there is nothing that the entoamoebas provoke to release serotonin, hence the serotonin concentration is very low. Low serotonin signals to the entoamoebas they are outside a host and they become less virulent to conserve energy. When they enter a new host, they multiply in the gut, and become more virulent as the enterochromaffine cells get provoked by them and the serotonin concentration increases.

text: Plants[edit]

text: In drying seeds, serotonin production is a way to get rid of the buildup of poisonous ammonia. The ammonia is collected and placed in the indole part of L-tryptophan, which is then decarboxylated by tryptophan decarboxylase to give tryptamine, which is then hydroxylated by a cytochrome P450 monooxygenase, yielding serotonin.[94]

text: However, since serotonin is a major gastrointestinal tract modulator, it may be produced by plants in fruits as a way of speeding the passage of seeds through the digestive tract, in the same way as many well-known seed and fruit associated laxatives. Serotonin is found in mushrooms, fruits and vegetables. The highest values of 25–400 mg/kg have been found in nuts of the walnut (Juglans) and hickory (Carya) genera. Serotonin concentrations of 3–30 mg/kg have been found in plantains, pineapples, banana, kiwifruit, plums, and tomatoes. Moderate levels from 0.1–3 mg/kg have been found in a wide range of tested vegetables.[17]

text: Serotonin is one compound of the poison contained in stinging nettles (Urtica dioica), where it causes pain on injection in the same manner as its presence in insect venoms (see below). It is also naturally found in Paramuricea clavata, or the Red Sea Fan.[95]

text: Serotonin and tryptophan have been found in chocolate with varying cocoa contents. The highest serotonin content (2.93 µg/g) was found in chocolate with 85% cocoa, and the highest tryptophan content (13.27–13.34 µg/g) was found in 70–85% cocoa. The intermediate in the synthesis from tryptophan to serotonin, 5-hydroxytryptophan, was not found.[96]

text: Invertebrates[edit]

text: Serotonin functions as a neurotransmitter in the nervous systems of most animals. For example, in the roundworm Caenorhabditis elegans, which feeds on bacteria, serotonin is released as a signal in response to positive events, such as finding a new source of food or in male animals finding a female with which to mate.[97] When a well-fed worm feels bacteria on its cuticle, dopamine is released, which slows it down; if it is starved, serotonin also is released, which slows the animal down further. This mechanism increases the amount of time animals spend in the presence of food.[98] The released serotonin activates the muscles used for feeding, while octopamine suppresses them.[99] Serotonin diffuses to serotonin-sensitive neurons, which control the animal's perception of nutrient availability.

text: If lobsters are injected with serotonin, they behave like dominant individuals whereas octopamine causes subordinate behavior.[21] A crayfish that is frightened may flip its tail to flee, and the effect of serotonin on this behavior depends largely on the animal's social status. Serotonin inhibits the fleeing reaction in subordinates, but enhances it in socially dominant or isolated individuals. The reason for this is social experience alters the proportion between serotonin receptors (5-HT receptors) that have opposing effects on the fight-or-flight response.[clarification needed] The effect of 5-HT1 receptors predominates in subordinate animals, while 5-HT2 receptors predominates in dominants.[100]

text: Insects[edit]

text: Serotonin is evolutionary conserved and appears across the animal kingdom. It is seen in insect processes in roles similar to in the human central nervous system, such as memory, appetite, sleep, and behavior.[101][18] Locust swarming is mediated by serotonin, by transforming social preference from aversion to a gregarious state that enables coherent groups.[102] Learning in flies and honeybees is affected by the presence of serotonin.[103][104] Insect 5-HT receptors have similar sequences to the vertebrate versions, but pharmacological differences have been seen. Invertebrate drug response has been far less characterized than mammalian pharmacology and the potential for species selective insecticides has been discussed.[105]

text: Wasps and hornets have serotonin in their venom,[106] which causes pain and inflammation.[15] as do scorpions.[107]

text: If flies are fed serotonin, they are more aggressive; flies depleted of serotonin still exhibit aggression, but they do so much less frequently.[108]

text: Growth and reproduction[edit]

text: In the nematode C. elegans, artificial depletion of serotonin or the increase of octopamine cues behavior typical of a low-food environment: C. elegans becomes more active, and mating and egg-laying are suppressed, while the opposite occurs if serotonin is increased or octopamine is decreased in this animal.[20] Serotonin is necessary for normal nematode male mating behavior,[109] and the inclination to leave food to search for a mate.[110] The serotonergic signaling used to adapt the worm's behaviour to fast changes in the environment affects insulin-like signaling and the TGF beta signaling pathway,[111] which control long-term adaption.

text: In the fruit fly insulin both regulates blood sugar as well as acting as a growth factor. Thus, in the fruit fly, serotonergic neurons regulate the adult body size by affecting insulin secretion.[112][113] Serotonin has also been identified as the trigger for swarm behavior in locusts.[114] In humans, though insulin regulates blood sugar and IGF regulates growth, serotonin controls the release of both hormones, modulating insulin release from the beta cells in the pancreas through serotonylation of GTPase signaling proteins.[24] Exposure to SSRIs during Pregnancy reduces fetal growth.[115]

text: Genetically altered C. elegans worms that lack serotonin have an increased reproductive lifespan, may become obese, and sometimes present with arrested development at a dormant larval state.[116][117]

text: Aging and age-related phenotypes[edit]

text: Serotonin is known to regulate aging, learning and memory. The first evidence comes from the study of longevity in C. elegans.[118] During early phase of aging[vague], the level of serotonin increases, which alters locomotory behaviors and associative memory.[119] The effect is restored by mutations and drugs (including mianserin and methiothepin) that inhibit serotonin receptors. The observation does not contradict with the notion that the serotonin level goes down in mammals and humans, which is typically seen in late but not early[vague] phase of aging.

text: Biochemical mechanisms[edit]

text: Biosynthesis[edit]

text: In animals including humans, serotonin is synthesized from the amino acid L-tryptophan by a short metabolic pathway consisting of two enzymes: tryptophan hydroxylase (TPH), aromatic amino acid decarboxylase (DDC) and the coenzyme pyridoxal phosphate. The TPH-mediated reaction is the rate-limiting step in the pathway. TPH has been shown to exist in two forms: TPH1, found in several tissues, and TPH2, which is a neuron-specific isoform.[120]

text: Serotonin can be synthesized from tryptophan in the lab using Aspergillus niger and Psilocybe coprophila as catalysts. The first phase to 5-hydroxytryptophan would require letting tryptophan sit in ethanol and water for 7 days, then mixing in enough HCl (or other acid) to bring the pH to 3, and then adding NaOH to make a pH of 13 for 1 hour. Asperigillus niger would be the catalyst for this first phase. The second phase to synthesizing tryptophan itself from the 5-hydroxytryptophan intermediate would require adding ethanol and water, and letting sit for 30 days this time. The next two steps would be the same as the first phase: adding HCl to make the pH = 3, and then adding NaOH to make the pH very basic at 13 for 1 hour. This phase uses the Psilocybe coprophila as the catalyst for the reaction.[121]

text: Serotonin taken orally does not pass into the serotonergic pathways of the central nervous system, because it does not cross the blood–brain barrier.[122] However, tryptophan and its metabolite 5-hydroxytryptophan (5-HTP), from which serotonin is synthesized, does cross the blood–brain barrier. These agents are available as dietary supplements, and may be effective serotonergic agents. One product of serotonin breakdown is 5-hydroxyindoleacetic acid (5-HIAA), which is excreted in the urine. Serotonin and 5-HIAA are sometimes produced in excess amounts by certain tumors or cancers, and levels of these substances may be measured in the urine to test for these tumors.

text: Effects of food content[edit]

text: Consuming purified tryptophan increases brain serotonin whereas eating foods containing tryptophan does not.[123] This is because the transport system which brings tryptophan across the blood-brain barrier is also selective for the other amino acids contained in protein sources.[122] High plasma levels of other large neutral amino acids compete for transport and prevent the elevated plasma tryptophan from increasing serotonin synthesis.

text: History and etymology[edit]

text: In 1935, Italian Vittorio Erspamer showed an extract from enterochromaffin cells made intestines contract. Some believed it contained adrenaline, but two years later, Erspamer was able to show it was a previously unknown amine, which he named "enteramine".[124] In 1948, Maurice M. Rapport, Arda Green, and Irvine Page of the Cleveland Clinic discovered a vasoconstrictor substance in blood serum, and since it was a serum agent affecting vascular tone, they named it serotonin.[125]

text: In 1952, enteramine was shown to be the same substance as serotonin, and as the broad range of physiological roles was elucidated, the abbreviation 5-HT of the proper chemical name 5-hydroxytryptamine became the preferred name in the pharmacological field.[126] Synonyms of serotonin include: 5-hydroxytriptamine, thrombotin, enteramin, substance DS, and 3-(β-Aminoethyl)-5-hydroxyindole.[127] In 1953, Betty Twarog and Page discovered serotonin in the central nervous system.[128]


document_title: List of Outlander episodes

question_text: how many episodes are there in outlander season 2

root

text: List of Outlander episodes

text: Series overview[edit]

text:
Season
Episodes
Originally aired

First aired
Last aired

1
16
8
August 9, 2014 (2014-08-09)
September 27, 2014 (2014-09-27)

8
April 4, 2015 (2015-04-04)
May 30, 2015 (2015-05-30)

2
13
April 9, 2016 (2016-04-09)
July 9, 2016 (2016-07-09)

3
13
September 10, 2017 (2017-09-10)
December 10, 2017 (2017-12-10)

tag: table

text: Episodes[edit]

text: Season 1 (2014–15)[edit]

text:
No.
overall
No. in
season
Title
Directed by
Written by
Original air date
U.S. viewers
(millions)

Volume One

1
1
"Sassenach"
John Dahl
Ronald D. Moore
August 9, 2014 (2014-08-09)
0.72[5]

In 1946, Claire Randall is a former WWII combat nurse visiting Inverness, Scotland, with her husband Frank, getting to know each other again after their wartime separation. Frank is researching his family history, in particular his 18th Century ancestor Jonathan "Black Jack" Randall. The morning after observing a modern Druid ritual at the standing stones on the hill of Craigh na Dun, Claire touches one of the stones, and wakes up lying on the ground. Soon she is in the middle of a skirmish between English redcoats and rebel Highlanders, and comes face to face with Frank's double, Randall, who is captain of the Dragoons. A Scotsman rescues Claire and she meets Jamie Fraser. She is able to set his dislocated arm and later treat his bullet wound, as they flee. Claire is brought to Castle Leoch, and realises that she has somehow travelled back in time.

2
2
"Castle Leoch"
John Dahl
Ronald D. Moore
August 16, 2014 (2014-08-16)
0.90[6]

Claire is presented to Colum MacKenzie, laird of the castle, and realises she is suspected of being an English spy. She meets Geillis Duncan, and witnesses Jamie doing a kindness for Laoghaire, the pretty young granddaughter of Mrs. Fitzgibbons. As Claire plans to return to Inverness and the standing stones, Colum decides it is best to keep her at Leoch, against her will, as healer to the clan.

3
3
"The Way Out"
Brian Kelly
Anne Kenney
August 23, 2014 (2014-08-23)
1.00[7]

Claire hopes that her medical skills will earn her the trust of the MacKenzies, and her eventual freedom, but manages only to make herself indispensable. By saving Mrs. Fitzgibbons's nephew from poisoning, which the fanatical Father Bain had declared as demonic possession, Claire makes a new enemy. After she and Jamie collude to help a youth punished for stealing, Claire is given renewed hope of being able to travel home when she hears a folk tale about a time traveller and the standing stones.

4
4
"The Gathering"
Brian Kelly
Matthew B. Roberts
August 30, 2014 (2014-08-30)
0.84[8]

Claire plots her escape during a clan gathering at Leoch, but Jamie convinces her that she would fail. Escorting her back, he finds himself in a dangerous position with Colum that he had been trying to avoid, but manages to make an oath of loyalty while manoeuvering himself out of trouble with his kinsman. Claire earns Dougal's respect while tending a dying man on a boar hunt, and he decides to take her with him on his rent collecting tour.

5
5
"Rent"
Brian Kelly
Toni Graphia
September 6, 2014 (2014-09-06)
0.95[9]

Claire clashes with Dougal and his men on the rent-collecting journey, especially when Dougal uses Jamie's scars to collect more money from the villagers. Later, she realises that they are collecting funds for the prospective Jacobite army. Knowing from history that their cause is doomed, she tries to warn them off.

6
6
"The Garrison Commander"
Brian Kelly
Ira Steven Behr
September 13, 2014 (2014-09-13)
1.10[10]

The redcoats take Claire to meet General Lord Thomas, who promises her safe passage to Inverness in the company of Randall. When the General leaves, Randall tries to discover why she is in Scotland. She concocts a story and he describes in detail his vicious whipping of young Jamie. Dougal comes to take her away, but Randall insists that he has to deliver Claire to Fort William the following day. They leave, and Dougal tells Claire he does not have to turn her in if she marries Jamie and so becomes Scottish.

7
7
"The Wedding"
Anna Foerster
Anne Kenney
September 20, 2014 (2014-09-20)
1.23[11]

Claire and Jamie get married to protect her from Randall, and try to get to know each other better afterwards to make the consummation less awkward. They realise their feelings for each other, but Claire is still torn thinking about Frank.

8
8
"Both Sides Now"
Anna Foerster
Ronald D. Moore
September 27, 2014 (2014-09-27)
1.42[12]

In 1946, Frank is losing hope of finding Claire, but goes to the standing stones one last time. In the past, Claire finds herself nearby to the stones, and runs for them. She hears Frank calling, and he hears her as well. But Claire is captured by English troops before she can reach the stones, and a despondent Frank leaves, as she is taken to Fort William. Claire nearly talks her way out of being interrogated by Randall, but he turns the tables on her. Just as he is about to rape her at knifepoint, Jamie appears in the window.

Volume Two

9
9
"The Reckoning"
Richard Clark
Matthew B. Roberts
April 4, 2015 (2015-04-04)
1.22[13]

Jamie saves Claire from Black Jack Randall, but the dynamic of their marriage is tested. Jamie tries to ease the conflict between Colum and Dougal sparked by Claire's rescue. Jamie rebuffs a seduction by Laoghaire, who then leaves a bad luck charm under Claire and Jamie's bed.

10
10
"By the Pricking of My Thumbs"
Richard Clark
Ira Steven Behr
April 11, 2015 (2015-04-11)
0.86[14]

Claire confronts a spiteful Laoghaire, but is surprised to learn that the charm came from Geillis. Jamie petitions the Duke of Sandringham for help to rid himself of Randall and the price on his head, while Claire secretly uses her historical knowledge to leverage his help in Jamie's favour. Dougal is inconsolable after his wife's death from an illness, but Geillis, pregnant with his child, is pleased. When Geillis's husband dies at a banquet for the Duke, Claire realises that Geillis poisoned him. Colum forbids Dougal from pursuing a relationship with Geillis, and exiles Dougal to his estate, with Jamie in tow. Claire and Geillis are arrested for witchcraft.

11
11
"The Devil's Mark"
Mike Barker
Toni Graphia
April 18, 2015 (2015-04-18)
1.09[15]

Claire and Geillis face charges of witchcraft. Ned Gowan comes to their defence, but witnesses, including Laoghaire and Father Bain, testify against them, and they are found guilty. Before Jamie can fight his way out of the courtroom with his wife, Geillis confesses and exonerates Claire. Claire recognises from the smallpox vaccine scar on Geillis's shoulder that she too is from the future. Geillis is taken to be burned at the stake. When Jamie asks Claire about her scar, she tells him of her time travel. He believes her, takes her back to Craigh na Dun, and wishes her well in returning to her own time. But Claire decides not to go back.

12
12
"Lallybroch"
Mike Barker
Anne Kenney
April 25, 2015 (2015-04-25)
1.11[16]

Jamie takes Claire to his family home of Lallybroch, where he clashes with his strong willed sister Jenny. Taking up his role as laird, he disrupts the household until Claire sets him straight. After Jamie narrowly avoids a run-in with passing redcoats, he and Jenny make amends.

13
13
"The Watch"
Metin Hüseyin
Toni Graphia
May 2, 2015 (2015-05-02)
1.05[17]

The men of the Watch arrive at Lallybroch, and Jamie pretends to be a Fraser cousin. Claire realises that Jenny's impending childbirth will be a dangerous, breech birth, but Jenny swears her to secrecy. The redcoat defector Horrocks appears, and threatens to reveal that Jamie has a price on his head unless Jamie pays him. Jenny's husband Ian kills Horrocks. Coming clean to the Watch leader MacQuarrie, Jamie agrees to join their raid. Jenny gives birth to a daughter with Claire's help. Meanwhile, the raiding party is ambushed, and Jamie captured by the redcoats.

14
14
"The Search"
Metin Hüseyin
Matthew B. Roberts
May 9, 2015 (2015-05-09)
1.08[18]

Claire and Jenny set out to find Jamie. They learn he has escaped his captors, and a newly arrived Murtagh takes Claire on a journey to find him. Their plan is to have Jamie find them by touring the countryside with Claire as a songstress called "The Sassenach". They attract Dougal's attention, and he tells them that Jamie has been recaptured and tried, and is awaiting execution at Wentworth Prison. Claire makes a deal with Dougal, and convinces his men to help her rescue Jamie.

15
15
"Wentworth Prison"
Anna Foerster
Ira Steven Behr
May 16, 2015 (2015-05-16)
1.01[19]

Randall appears at Wentworth Prison and tortures Jamie, seeking his submission. Meanwhile, Claire gets into the prison to save her husband. When Randall catches her, Jamie agrees to submit to rape by Randall in exchange for her freedom. Before she leaves, Claire "curses" Randall and tells him what will be the date of his death. When all seems lost, Murtagh comes up with a plan to rescue Jamie.

16
16
"To Ransom a Man's Soul"
Anna Foerster
Ira Steven Behr & Ronald D. Moore
May 30, 2015 (2015-05-30)
0.98[20]

Murtagh's rescue is successful and Randall left for dead, but Jamie remains tortured by his ordeal. As Claire tends to his injuries, Jamie is tormented by memory of his rape and torture. Murtagh accompanies them as they set sail for safety in France, where Claire hopes to meet Charles Stuart and somehow prevent the catastrophic Battle of Culloden. As they set sail, Claire tells Jamie that she is pregnant.

text: Season 2 (2016)[edit]

text:
No.
overall
No. in
season
Title
Directed by
Written by
Original air date
U.S. viewers
(millions)

17
1
"Through a Glass, Darkly"
Metin Hüseyin
Ronald D. Moore
April 9, 2016 (2016-04-09)
1.46[21]

Claire awakes at Craigh Na Dun back in 1948. Grieving over Jamie's assumed death in battle, she is reunited with Frank, who cannot understand her reappearance but is desperate to reconcile. Claire explains where she has been, including her relationship with Jamie and that she is carrying his child. They agree that they will raise the child together and resume married life. She later recalls her arrival with Jamie to France, where they make contact with Jamie's Jacobite cousin Jared Fraser, and make an enemy of the Comte St. Germain.

18
2
"Not in Scotland Anymore"
Metin Hüseyin
Ira Steven Behr
April 16, 2016 (2016-04-16)
1.24[22]

In 1744, Claire and Jamie have settled in Paris, ostensibly working for Jared but actually seeking to foil the Jacobite rebellion. Jared arranges for Jamie to meet with Prince Charles Stuart himself, but Jamie cannot deter him from planning to seize the English throne for his father. Claire secures an invitation to the Palace of Versailles, where she hopes to meet King Louis XV of France and thwart his funding of the Stuart cause. The Frasers make an impression, and are reunited with the Duke of Sandringham. Claire is shocked to learn that Randall is still alive, and is not sure if she should tell Jamie.

19
3
"Useful Occupations and Deceptions"
Metin Hüseyin
Anne Kenney
April 23, 2016 (2016-04-23)
1.21[23]

While Jamie continues to try and sour the financing of Prince Charles's rebellion, Claire makes herself useful at L'Hôpital des Anges, a charity hospital run by the imperious Mother Hildegarde. Claire and Jamie discover that Sandringham is helping Charles secure funds from English nobles, which Jamie hopes to thwart. Though spending time with Sandringham will surely reveal to Jamie that Randall is alive, Claire cannot bear to tell Jamie herself.

20
4
"La Dame Blanche"
Douglas Mackinnon
Toni Graphia
April 30, 2016 (2016-04-30)
1.17[24]

Claire finally tells Jamie about Randall, and the potential for vengeance reinvigorates Jamie. Claire suspects the Comte St. Germain when an attempt is made on her life, and she and Jamie plan a dinner party to paint Charles and his cause in a bad light to Sandringham. Murtagh, Claire and the young Mary Hawkins are attacked on their way to the party, and Mary is raped. Back at the house, the party goes as planned until a misunderstanding sparks a brawl.

21
5
"Untimely Resurrection"
Douglas Mackinnon
Richard Kahan
May 7, 2016 (2016-05-07)
1.13[25]

The disastrous dinner party has discouraged Charles's investors, but the prince finds a new one in Jamie's enemy, St. Germain. Charles puts Jamie and St. Germain into business together, to their concern. Trying to preserve Frank's lineage, Claire sabotages the blossoming relationship between Mary Hawkins and Alex Randall. Randall appears at Versailles, and Jamie challenges him to a duel. Jamie is angry when Claire forces him to swear that he will not harm Randall until he has conceived a child with Mary.

22
6
"Best Laid Schemes..."
Metin Hüseyin
Matthew B. Roberts
May 14, 2016 (2016-05-14)
1.03[26]

Jamie tells Murtagh the truth about Claire's past. Claire uses her medical knowledge to give St. Germain's men symptoms that resemble those of smallpox. Jamie and Murtagh then steal the Comte's wine shipment, seemingly dashing Charles's hopes. Fate brings Jamie and Randall face to face, and Claire arrives just in time to see them duelling. As Jamie stabs Randall and gendarmes arrive to arrest them, a pregnant Claire collapses, bleeding.

23
7
"Faith"
Metin Hüseyin
Toni Graphia
May 21, 2016 (2016-05-21)
1.10[27]

Claire and Jamie's daughter is stillborn, and a devastated Claire nearly dies from fever. With Jamie in the Bastille, Claire discovers that he duelled with Randall because he had discovered the English captain raping Fergus. Claire seeks mercy for Jamie from the king, who forces her to decide the fates of suspected "sorcerers" Master Raymond and St. Germain. She contrives to save them, but Raymond poisons St. Germain. Claire submits sexually to the king in exchange for Jamie's freedom. They reconcile, and visit their daughter Faith's grave before leaving Paris for Scotland.

24
8
"The Fox's Lair"
Mike Barker
Anne Kenney
May 28, 2016 (2016-05-28)
1.06[28]

In Scotland, Jamie finds himself compelled to support the Jacobite rebellion. He and Claire seek military assistance from his grandfather, Lord Lovat, and are reunited with Colum and Laoghaire. Colum hopes to discourage the rebellion, which he is certain will fail. Laoghaire begs Claire's forgiveness, and later agrees to help Claire and Jamie manipulate Lovat. Playing both sides, Lovat promises peace to Colum, but also sends Jamie some troops as reinforcements.

25
9
"Je Suis Prest"
Philip John
Matthew B. Roberts
June 4, 2016 (2016-06-04)
0.94[29]

Claire and Jamie are reunited with Angus, Rupert and Dougal. As Jamie and Murtagh attempt to train Lovat's men, the camps remind Claire of her traumatic experiences in World War II. She and Jamie later coerce young English scout William Grey to reveal vital information about the English army that is nearby.

26
10
"Prestonpans"
Philip John
Ira Steven Behr
June 11, 2016 (2016-06-11)
0.82[30]

Jamie and the Jacobite army undertake their first battle against the English at Prestonpans. Surprising them in the fog, the Highlanders are victorious, but Angus later dies of his injuries.

27
11
"Vengeance Is Mine"
Mike Barker
Diana Gabaldon
June 18, 2016 (2016-06-18)
0.86[31]

When Claire, Jamie and their party are surrounded by redcoats, Claire pretends to be an English hostage so Dougal can negotiate the Highlanders' safe passage. Taken to 'safety', Claire finds herself face to face with the Duke of Sandringham and his goddaughter Mary Hawkins. She also recognises Sandringham's valet, Danton, as the man who had raped Mary. Sandringham intends to lure "Red Jamie" to save Claire, and then turn them both over to the redcoats. Forewarned, Jamie and Murtagh slip into the castle. Mary kills Danton, and Murtagh beheads Sandringham to fulfill his promise to Claire and Mary to avenge their attack in Paris.

28
12
"The Hail Mary"
Philip John
Ira Steven Behr & Anne Kenney
June 25, 2016 (2016-06-25)
1.05[32]

Jamie tries to avoid the Battle of Culloden with a preemptive strike against the English while they are celebrating the Duke of Cumberland's birthday, but the Prince gets lost in the dark. Claire finds herself trying to convince Randall to marry Mary, who is pregnant with a dying Alex's child. A dying Colum asks Claire to help him end his life painlessly. Before she does, he tells her that Geillis was not burned at the stake until after her child with Dougal was born, and that the child was adopted into the MacKenzie clan.

29
13
"Dragonfly in Amber"
Philip John
Toni Graphia & Matthew B. Roberts
July 9, 2016 (2016-07-09)
1.15[33]

In 1968, fifty-year-old Claire is reunited with Reverend Wakefield's adopted son, Roger, in Inverness. Her twenty-year-old daughter, Brianna, discovers that she is not Frank's biological daughter, but refuses to believe her mother's tale of time travel. Meanwhile, Brianna meets Scottish Independence activist Gillian Edgars, whom Claire later recognises is Geillis Duncan. Before the battle of Culloden in 1746, Jamie insists that Claire save herself and their unborn child by travelling back through the stones. Back in 1968, Claire realises Roger is a descendant of Geillis and Dougal. To prevent Gillian's death, they try to stop her from travelling back through time. They arrive too late, but seeing Gillian disappear into the stone convinces Brianna that her mother is telling the truth. Roger reveals to Claire that Jamie did not die at Culloden.

text: Season 3 (2017)[edit]

text:
No.
overall
No. in
season
Title
Directed by
Written by
Original air date
U.S. viewers
(millions)

30
1
"The Battle Joined"
Brendan Maher
Ronald D. Moore
September 10, 2017 (2017-09-10)
1.49[34]

In 1948, a pregnant Claire moves to Boston with Frank. Their relationship is tense, and she struggles to adjust to her new life as a housewife. She gives birth to Jamie's daughter. In 1746, Jamie and Rupert have survived the Battle of Culloden, but Jamie's wounds are grave. They are caught by redcoats, and Rupert is executed. Just as Jamie's turn comes, he is recognised by Lord Melton, the elder brother of the boy—John Grey—whose life Jamie spared before Prestonpans. Bound by honour, Melton sends Jamie home to Broch Tuarach, expecting him to die en route.

31
2
"Surrender"
Jennifer Getzinger
Anne Kenney
September 17, 2017 (2017-09-17)
1.40[35]

Baby Brianna has brought Claire and Frank closer, but the memory of Jamie stands between them. Searching for fulfillment, Claire enrolls in medical school as the only woman in her class. Jamie has survived, but with the English searching for him, he lives in a cave as an outlaw. Soldiers come to Lallybroch repeatedly, harassing Jenny and Ian for information about Jamie's whereabouts. When Fergus loses a hand in a run-in with redcoats, Jamie arranges to be "caught", so that Jenny can claim the reward and the English will leave his family alone.

32
3
"All Debts Paid"
Brendan Maher
Matthew B. Roberts
September 24, 2017 (2017-09-24)
1.55[36]

Claire graduates. Years later as Brianna graduates high school, Frank tells Claire he wants a divorce so he can marry his mistress and return to England. They are poised to argue over where Brianna will live, but Frank is killed in a car accident. Jamie is serving time in Ardsmuir Prison, where he is the leader and spokesman for the Scottish prisoners. His uneasy relationship with the new governor, an adult John Grey, evolves into friendship. When the prison is shuttered, Murtagh and the rest of the prisoners are sent to the American colonies as indentured servants, while Grey paroles Jamie to work at a private estate called Helwater.

33
4
"Of Lost Things"
Brendan Maher
Toni Graphia
October 1, 2017 (2017-10-01)
1.59[37]

Claire and Brianna research history to track Jamie to Ardsmuir, but when the trail runs cold, they return to America. Serving as a groom for the Dunsany family, Jamie is blackmailed into sex by heiress Geneva. When she and her earl husband die, leaving their newborn son Willie an orphan, Geneva's sister Isobel keeps the secret that the child is really Jamie's. Years later, when the young earl's resemblance to him is becoming obvious, Jamie leaves Helwater, with Willie in the care of Isobel and Grey, who are engaged to be married.

34
5
"Freedom & Whisky"
Brendan Maher
Toni Graphia
October 8, 2017 (2017-10-08)
1.60[38]

In December 1968, Roger arrives in Boston having discovered a clue to Jamie's whereabouts in 1765. Brianna urges Claire to go back and reunite with Jamie, and mother and daughter have a tearful farewell. In Edinburgh in 1765, Claire finds Jamie in a print shop.

35
6
"A. Malcolm"
Norma Bailey
Matthew B. Roberts
October 22, 2017 (2017-10-22)
1.72[39]

Claire and Jamie are reunited. She shares photos of Brianna, and he tells Claire about Willie. She also learns that Jamie prints subversive material, smuggles wine and spirits, and lives in a brothel. Claire is reunited with an adult Fergus, and meets some of the prostitutes in Madame Jeanne's establishment. While Jamie is out, Claire finds herself in danger from an intruder.

36
7
"Crème de Menthe"
Norma Bailey
Karen Campbell
October 29, 2017 (2017-10-29)
1.52[40]

Claire's attacker falls and suffers a critical head injury, and she is unable to save his life. Ian arrives looking for his son, Young Ian. Jamie claims he has not seen the boy, though young Ian has been working for him. Young Ian confronts an intruder at the print shop, who discovers Jamie's treason. The shop is soon consumed by fire, but Jamie saves the boy.

37
8
"First Wife"
Jennifer Getzinger
Joy Blake
November 5, 2017 (2017-11-05)
1.64[41]

Jamie takes Claire and young Ian home to Lallybroch. Claire discovers that Jamie married a twice widowed Laoghaire, who accidentally shoots him during a confrontation. Jamie recovers thanks to the penicillin that Claire has brought. Ned Gowan negotiates a settlement with Laoghaire, and Jamie decides to retrieve the treasure on Silky Isle to pay for it. Young Ian swims to the island in an injured Jamie's place. Jamie and Claire are horrified to see the boy forcibly taken aboard a ship and carried away.

38
9
"The Doldrums"
David Moore
Shannon Goss
November 12, 2017 (2017-11-12)
1.49[42]

Learning in France that young Ian's ship is heading for Jamaica, Claire and Jamie pursue it on a trading ship, the Artemis. At sea, Fergus reveals that Laoghaire's daughter Marsali is aboard, and they seek Jamie's approval for their marriage. The British man-of-war Porpoise stops them and seeks help, her crew stricken with a plague. Claire determines that the disease is typhoid fever and trains them to stop its spread, but the ship carries her off with the captain's promise to leave her in Jamaica to wait for the Artemis.

39
10
"Heaven and Earth"
David Moore
Luke Schelhaas
November 19, 2017 (2017-11-19)
1.23[43]

On the Porpoise, Claire struggles against suspicion from some of the crew, but eventually manages to stop the spread of the disease. She discovers that the captain knows Jamie's true identity and plans to have him arrested in Jamaica for treason and murder. When they go ashore, she tries to escape but is apprehended by the captain. That night at sea, a woman named Annejke encourages Claire to jump overboard using an empty barrel as a raft, as the current will carry her towards land. Meanwhile on the Artemis, Jamie is confined after threatening the captain, wanting him to sail faster in pursuit of the Porpoise. He tries to get Fergus to help him take control of the ship, but Fergus instead persuades the captain to let Jamie rejoin the crew after he promises not to mutiny again.

40
11
"Uncharted"
Charlotte Brandström
Karen Campbell & Shannon Goss
November 26, 2017 (2017-11-26)
1.40[44]

Claire swims ashore, and walks inland, struggling to survive the harsh tropical environment. She is found and taken in by Father Fogden, a runaway priest. In passing, he tells her of a cave in Jamaica where people have been known to disappear. A few days later, the Artemis reaches the island, needing to make repairs after a storm, her captain having died. As the ship is about to sail, Claire attracts Jamie's attention from the beach using a mirror. Back on the island, Fogden marries Fergus and Marsali. The next day, Jamie, Claire and the crew set sail for Jamaica.

41
12
"The Bakra"
Charlotte Brandström
Luke Schelhaas
December 3, 2017 (2017-12-03)
1.51[45]

Young Ian is being held prisoner in Jamaica by Geillis. She makes him drink tea that compels him to tell the truth about Jamie and the treasure. Jamie and Claire reach the island, and Claire gets into an altercation at the slave market. Jamie buys her the slave, Temeraire, to defuse the situation. They attend a ball thrown by the new governor, who they discover is Lord John. Claire is reunited with Geillis, who offers to help with their search. At the ball, Margaret Campbell prophesies that a king will rule Scotland again when a two-hundred year-old child is killed. Fergus sees Captain Leonard arrive, and warns Jamie, who flees. Temeraire tells Claire and Jamie that Geillis is holding Ian. On their way to Geillis's villa, Jamie is arrested by Leonard.

42
13
"Eye of the Storm"
Matthew B. Roberts
Matthew B. Roberts & Toni Graphia
December 10, 2017 (2017-12-10)
1.43[46]

Claire is captured whilst searching Geillis's slave houses for Ian. The governor's troops recover Jamie from Captain Leonard, and Grey sets Jamie free after Leonard is unable to produce any evidence of his alleged crimes. Meanwhile, Claire tells Geillis about her return to the 20th century, and Geillis realises that it is Claire's child who must die before Scotland wins its new king. Jamie frees Claire, and they rescue Ian from Geillis, who is killed during the struggle. The group sets sail from Jamaica, but are shipwrecked in a storm. Jamie saves Claire from drowning, and they are washed up together on the shore of Georgia.

text: Ratings[edit]

text:
Season
Ep. 1
Ep. 2
Ep. 3
Ep. 4
Ep. 5
Ep. 6
Ep. 7
Ep. 8
Ep. 9
Ep. 10
Ep. 11
Ep. 12
Ep. 13
Ep. 14
Ep. 15
Ep. 16

1
0.72
0.90
1.00
0.84
0.95
1.10
1.23
1.42
1.22
0.86
1.09
1.11
1.05
1.08
1.01
0.98

2
1.46
1.24
1.21
1.17
1.13
1.03
1.10
1.06
0.94
0.82
0.86
1.05
1.15
N/A

3
1.49
1.40
1.55
1.59
1.60
1.72
1.52
1.64
1.49
1.23
1.40
1.51
1.43
N/A


document_title: Fortitude (TV series)

question_text: where is the town of fortitude supposed to be

text: Fortitude (TV series)

text: Production[edit]

text: Conception[edit]

text: Writer Simon Donald based his parasite on ichneumon wasps, which lay their eggs in hosts, after talking to a parasitologist.[6]

tag: p

text: Location[edit]

text: Fortitude is a fictional community located on Svalbard in Arctic Norway. It is described as an international community, with inhabitants from many parts of the world (population of 713 inhabitants and 4 police officers).[7] The series was filmed in both the UK and in Reyðarfjörður, Iceland.[8][9]

text: Casting[edit]

text: American actor Stanley Tucci appears in his first British television role as Detective Chief Inspector Morton from London's Metropolitan Police, who has flown to the peaceful close-knit community of Fortitude, as the local sheriff, played by Richard Dormer, investigates a violent murder. Christopher Eccleston plays Charlie Stoddart, a British scientist who leads the arctic biology department at the Fortitude arctic research facility. Michael Gambon portrays Henry Tyson, a wildlife photographer who is dying of cancer.[10] The Killing star Sofie Gråbøl plays Hildur Odegard, the governor of Fortitude, in her first UK television drama role.[11][12]

text: Cast[edit]

text: Main cast[edit]

text:
Richard Dormer as Police Station Chief/Sheriff Dan Anderssen
Sofie Gråbøl as Governor Hildur Odegard
Björn Hlynur Haraldsson as Police Sergeant Eric Odegard
Mia Jexen as Police Constable Ingrid Witrey
Alexandra Moen as Police Constable Petra Bergen
Sienna Guillory as Natalie Yelburton
Luke Treadaway as Vincent Rattrey
Darren Boyd as Markus Huseklepp
Verónica Echegui as Elena Ledesma
Ramon Tikaram as Tavrani Tavra
Nicholas Pinnock as Frank Sutter (Series 1)
Jessica Raine as Julia 'Jules' Sutter (Series 1)
Christopher Eccleston as Professor Charlie Stoddart (Series 1)
Stanley Tucci as Detective Chief Inspector Eugene Morton (Series 1)
Michael Gambon as Henry Tyson (Series 1)
Jessica Gunning as Shirley Allerdyce (Series 1)
Johnny Harris as Ronnie Morgan (Series 1)
Aaron McCusker as Jason Donnelly (Series 1)
Phoebe Nicholls as Dr. Margaret Allerdyce (Series 1)
Dennis Quaid as Michael Lennox (Series 2)
Michelle Fairley as Freya Lennox (Series 2)[13]
Edvin Endre as Rune Lennox (Series 2)
Ken Stott as Erling Munk (Series 2)
Parminder Nagra as Dr. Surinder Khatri (Series 2)[14]
Robert Sheehan as Vladek Klimov (Series 2)
Brian McCardie as Lazlo Hindemith (Series 2)
Paul Ready as Mark Devlin (Series 2)

tag: ul

text: Recurring cast[edit]

text: Series 1[edit]

text:
Chipo Chung as Trish Stoddart
Darwin Brokenbro as Liam Sutter
Elizabeth Dormer-Phillips as Carrie Morgan
Michael Obiora as Max Cordero
Emil Hostina as Yuri Lubimov
Jonjo O'Neill as Ciaran Donnelly
Leanne Best as Celia Donnelly
Lorcan Cranitch as DI Bernard Littlejohn

text: Series 2[edit]

text:
Jude Akuwudike as Dr. Adebimpe
Gershwyn Eustache, Jnr. as Tomak Bowzyk
Rune Temte as Lars Ulvinaune
Eloise Smyth as Yeva Podnikov
David Yip as Hong Mankyo
Ralph Riach as Ralfi Sigurdson
Joi Johanssen as Axel Fersen
Steve Toussaint as Avec Lamont
Mark Field as 'The Fiend'
Jessica Henwick as Bianca Mankyo

text: Episodes[edit]

text: Series 1 (2015)[edit]

text:
No.
overall
No. in
series
Title
Directed by
Written by
Original air date
UK viewers
(millions) [15]

1
1
"Episode 1"
Sam Miller
Simon Donald
29 January 2015 (2015-01-29)
1.53

Professor Charlie Stoddart, a scientist at the Fortitude research centre, is approached by Jason Donnelly, who claims to have found a mammoth carcass. When Jason offers to sell Charlie the carcass, he refuses and threatens to report Donnelly to the police for the illegal sale of a scientific discovery. Meanwhile, a young boy, Liam Sutter, faints after taking a walk with his friend, Carrie Morgan. When he later awakens, he discovers he is alone. His father, Frank, who is supposed to be looking after him, is nowhere to be seen. Liam opens up his bedroom window, and climbs barefoot into the snow. When he develops frostbite, he is rushed to hospital. Meanwhile, scientist Vincent Rattrey arrives at Stoddart's house for a pre-planned meeting and finds him dead, the victim of a savage attack.

2
2
"Episode 2"
Sam Miller
Simon Donald
29 January 2015 (2015-01-29)
1.53

Rattrey is taken to the police station, but as forensic officers arrive to sweep the crime scene, they discover Stoddart is still alive, but he dies on the way to hospital. The mysterious disappearance of Natalie Yelburton leads Sheriff Anderssen to suspect she is either the perpetrator or another victim. When Henry Tyson discovers Stoddart has been murdered, he makes a call to an old friend in Britain. DCI Eugene Morton, from the London Metropolitan Police, arrives to investigate. Anderssen isn't pleased at Morton's arrival and immediately makes his feelings known. Dr. Allerdyce offers Liam's mother Julia the chance for Liam to take part in an experimental procedure which could stave the effects of frostbite.

3
3
"Episode 3"
Sam Miller
Simon Donald
5 February 2015 (2015-02-05)
1.26

The search for Natalie Yelburton continues, but information from Eric Odegard points the team toward Jason Donnelly, who they suspect may be responsible for Stoddart's murder, who he confronted earlier in the day. Sheriff Anderssen is suspicious when he learns Morton left London before Stoddard's murder was even reported. Meanwhile, Morton receives advice from local teacher Markus Huseklepp, who turns to him after a violent confrontation with Frank Sutter. As the search for Natalie and Jason leads the team to an exploration bunker out on the glacier, they finally stumble upon a possible motive for the murder. As the team set out to find Jason's brother Ciaran, Governor Hildur discovers Stoddart left a telephone message for his wife, which could thwart her plans to build a hotel on the glacier.

4
4
"Episode 4"
Richard Laxton
Stephen Brady
12 February 2015 (2015-02-12)
1.10

Ciaran is found beaten and gagged in a warehouse at the back of his shop, and he points the finger towards Ronnie Morgan. Morgan is nowhere to be found, and his daughter, Carrie, is also missing. Meanwhile, Morton seizes his chance to interview Henry Tyson, who denies making the phone call that brought Morton to Fortitude. When Eric Odegard pays a visit to Julia, an unexpected accident leads him to discover Frank's blood-stained T-shirt in the bathroom. Eric tries to alert Sheriff Anderssen, but he is out of radio range while searching for a boat used by Ronnie Morgan in his escape. Later, a vital piece of evidence belonging to Stoddart is stolen from Morton's hotel room while he is distracted by an altercation next door, where he discovers an angry Sheriff Anderssen savagely beating Frank.

5
5
"Episode 5"
Richard Laxton
Simon Donald
19 February 2015 (2015-02-19)
1.15

Frank is taken in and interviewed by Morton. He denies being responsible for Stoddard's murder and claims, on the night in question, he found his son covered in blood, saying this caused by a ruptured swelling in his son's throat. Hildur organises a DNA test to confirm if the blood found on the shirt is Stoddard's or Liam's. Elena reveals he snuck out to have sex with her and left his son alone. The blood is confirmed to belong to Stoddard, but Frank is adamant that he found his son covered in blood and was not responsible for the murder. When Morton finds a pair of blood-stained pyjamas hidden in Liam's room, the evidence seemingly confirms Frank's story. Meanwhile, Ronnie Morgan continues to lay low from the police, camped out on the glacier with his daughter.

6
6
"Episode 6"
Hettie MacDonald
Simon Donald
26 February 2015 (2015-02-26)
1.15

In a flashback to the night of the murder, Liam is seen leaving his bedroom and walking over to Stoddard's house. He attacks him with a chopping board before taking cover under the kitchen table. Further forensic examination of Stoddard's body reveals a broken fingernail lodged in a rib, later determined to belong to Liam. Dr. Allerdyce wakens Liam in his hyperbaric chamber just long enough for him to confirm to Morton he was present at the murder scene. Jason discovers Ronnie has taken the ivory tusks from the Mammoth carcass. Meanwhile, the focus of Morton's investigation switches to the dead Pettigrew, a geologist who was eaten by a polar bear. After discovering who was responsible for stealing the evidence, he unexpectedly stumbles upon a possible crime scene.

7
7
"Episode 7"
Hettie MacDonald
Tom Butterworth & Chris Hurford
5 March 2015 (2015-03-05)
1.05

As Fortitude begins to come to terms with Stoddard's death, Morton begins his investigation to find out who was responsible for Pettigrew's murder. After visiting the research centre and speaking with lab technician Max Cordero, he decides to track down Yuri Lubimov, whom he is certain holds vital information on the case. Shirley Allerdyce becomes more and more sick, but her boyfriend Markus continues his quest to make her 'more beautiful by the day'. Lubimov suggests to Morton that Eric was responsible for Pettigrew's murder, and that his police colleagues have hidden evidence in order to protect him. Henry Tyson has a flashback and realises he photographed Pettigrew's murder scene. Ronnie Morgan arranges a deal to sell the ivory tusks but is ambushed and is forced to hand over his goods at gunpoint. Shirley's illness overcomes her, leading her to brutally attack her own mother and leave her for dead.

8
8
"Episode 8"
Hettie MacDonald
Ben Richards
12 March 2015 (2015-03-12)
1.00

The townsfolk are disturbed by the second attack and suspect a serial killer may be on the loose. Shirley is found dead of a heart attack in a nearby supermarket. Frank Sutter becomes convinced the second attack proves his son was not responsible for Stoddart's murder and tries to find evidence to prove Markus' involvement. As Dr. Allerdyce fights for her life in hospital, Carrie Morgan finally returns home.

9
9
"Episode 9"
Nick Hurran
Stephen Brady
19 March 2015 (2015-03-19)
1.03

Frank Sutter interrogates Huseklepp, convinced he was responsible for his son's involvement in Stoddart's death. Meanwhile, Rattrey and Yelburton receive a note from Stoddard, to be delivered in the event of his death. It asks them to perform a brain analysis on a polar bear who was found to have mauled and eaten another adult male. Toxins in the bear's brain motivate Rattrey to perform the same analysis on Shirley Allerdyce to determine if her body carried the same toxins. When the test results come back positive, Rattrey realises an environmental factor played a part in both murders. Morton continues to probe into the circumstances surrounding Pettigrew's death. When he finds a series of photographs in Henry Tyson's dark room depicting the scene, he realises Sheriff Anderssen's version of events doesn't tally up.

10
10
"Episode 10"
Nick Hurran
Tom Butterworth & Chris Hurford
26 March 2015 (2015-03-26)
0.87

The test to determine if Liam Sutter carries the same toxins as Shirley Allerdyce comes back as negative, forcing Rattrey and Yelburton to seek approach. They discover both perpetrators carried spores in their blood, suggesting an airborne virus. When Jules decides to flee with Liam, they discover Hildur locked down the island in order to determine who has been affected. Jason Donnelly returns home, battered and bruised, but refuses to give an account of his whereabouts. Morton heads out onto the glacier in search of Henry, but meets a grisly fate when Henry refuses to be interrogated by Morton. Huseklepp becomes increasingly anxious when the research center refuse him access to Shirley's body.

11
11
"Episode 11"
Sam Miller
Simon Donald
2 April 2015 (2015-04-02)
0.82

Anderssen heads out onto the glacier in an attempt to rescue Morton. In a flashback to the night of Pettigrew's death, Anderssen reveals to Morton that he found Pettigrew in Elena's hotel room after raping her. After subduing him, he took Pettigrew to the beach and handcuffed him to a pylon, where he was subsequently mauled by a polar bear before being shot by Henry Tyson. Meanwhile, Rattrey goes in search of Stoddart's dog, shot at the scene of his murder. A post-mortem examination reveals the dog ingested larvae found in Stoddart's remains. Suspecting Dr. Allerdyce became the larvae's host, transferring them from Liam Sutter to her daughter, he finally uncovers the link between the two murders. But when attempting to uncover the larvae's identity, he finds Dr. Allerdyce's body has become a feeding ground.

12
12
"Episode 12"
Sam Miller
Simon Donald
9 April 2015 (2015-04-09)
1.06

Rattrey is trapped with Dr. Allerdyce in her hospital room, forced to succumb to the plague of wasps, as Yelburton and Sheriff Anderssen watch helplessly. Meanwhile, Eric goes out in search of the missing ice drill, and finds Yuri and Max out on the glacier. Yuri, powered by his own greed, decides to fight Eric, leaving him unconscious – but he himself is left helpless when he falls into a cave beneath the ice, into a mammoth's graveyard. Jason finally reveals the location of the mammoth carcass, and the location of Ronnie Morgan's body – but it all seems too late. As Elena succumbs to the virus, Anderssen is forced to shoot her to prevent her from infecting the child.

text: Series 2 (2017)[edit]

text:
No.
overall
No. in
series
Title
Directed by
Written by
Original air date
UK viewers
(millions) [15]

13
1
"Episode 1"
Hettie MacDonald
Simon Donald
26 January 2017 (2017-01-26)
0.777

A road traffic accident out on the glacier leads to the discovery of a headless body. Forensic examination reveals that the angle of decapitation suggests that it could not have been an accident. As Ingrid and Petra try to identity the victim, Hildur is forced to take charge of the investigation in Anderssen and Eric's absence. Meanwhile, fisherman Michael Lennox has pitched up in Fortitude in the hope of collecting an incoming haul of rare Arctic King Crab worth upward of $13 per pound. However, a fire on his vessel leads him to suspect that one of his rivals may have discovered his intended plan. Vincent announces that he is heading back home to London for good, but not before he reveals his true feelings for Natalie. Convinced that the former sheriff has died somewhere out on the glacier, Hildur sends out a search party to bring Eric home, but then Anderssen unexpectedly reappears.

14
2
"Episode 2"
Hettie MacDonald
Tom Butterworth & Chris Hurford
2 February 2017 (2017-02-02)
0.350

Hildur is shocked to discover that she is to be replaced as governor of Fortitude by Munk. The search for the glacier victim's missing head takes a surprising turn when Ingrid discovers an item of clothing belonging to Rune in the victim's house. Attempting to use Tavrani as a source of information to discover how the decapitation may have taken place, Eric is surprised to find the victim's missing head upright in Tavrani's freezer. Tavrani manages to escape, leaving Eric and the team no closer to uncovering the killer's identity. Meanwhile, Michael and Markus are forced to join forces when a polar bear runs rampage in the local primary school, leaving Vincent and Natalie attempting to identify the cause of the animal's rage. As Hildur tries to discover why her bosses are keen to oust her out, Anderssen continues to fight for his life.

15
3
"Episode 3"
Hettie MacDonald
Simon Burke
9 February 2017 (2017-02-09)
0.330

Eric, Ingrid and Petra finally manage to track down Tavrani, but he fails to provide them with any useful information as to who the killer might be. Meanwhile, Natalie and Vincent head out into the wilderness in an attempt to discover why a deer has been spotted eating the remains of a polar bear in the frigid wastes of the tundra. When a second headless victim is found out on the glacier, Munk angrily demands Eric to catch the killer. However, the killer's reign of terror continues to cause trouble when a burial site is savaged for surviving organs. Whilst trapped in custody, a masked intruder forces his way into the station and uses a knife to remove Tavrani's tongue. Khatri orders a review of Anderssen's medical history in light of his miraculous recovery. Hildur struggles to adjust to life outside of government.

16
4
"Episode 4"
Kieron Hawkes
Simon Burke
16 February 2017 (2017-02-16)
0.298

Hong Mankyo identifies the second headless victim found out on the glacier as his missing daughter, Bianca. Tavrani begins to recover in hospital, but continues to refuse to identify who was responsible for the vicious attack. Hildur and Eric's relationship continues to crumble as it becomes apparent that Munk has ulterior motives for becoming acting governor. Michael offers Hildur assistance in retrieving communications from Munk's computer that could provide information as to why she was sacked. Vladek Klimov comes to Fortitude and reignites his friendship with Freya, Michael Lennox's wife, and she asks Vladek, a shaman, to heal her from her deadly neuro-degenerative disease (amyotrophic lateral sclerosis, AKA Lou Gehrig's disease). Natalie and Vincent's field trip continues to spiral when Vincent begins to suffer from a bout of hysterical blindness. Eric comes face to face with Anderssen for the first time since his miraculous recovery.

17
5
"Episode 5"
Kieron Hawkes
Simon Donald
23 February 2017 (2017-02-23)
0.168

Just hours after his release from hospital, Tavrani is found dead of suspected heart failure. Later, during the memorial service for Bianca Mankyo, a hooded figure disrupts the mourners by sadistically taunting them with Tavrani's missing tongue. Eric and Michael give chase and are led into a mysterious lair where they find Bianca's missing head suspended from the ceiling in a yellow holdall. The man escapes, but not before Eric shoots him. Hildur is left a suspicious package in her hotel room, which contains video footage of an autopsy from 1942 in which the victim appears to return to life. Freya tends to Vladek Klimov's wound, revealing him to be the man Eric shot. Anderssen threatens Khatri to gain entry into the research facility to turn off Elena's life support machine, but is shocked when she makes an unexpected recovery. Ingrid suspects that her mother has been harbouring Vladek Klimov, the man suspected of being the murderer. Munk warns bartender Tomak that he is no longer prepared to support his activities, revealing the two to be conspiring.

18
6
"Episode 6"
Metin Hüseyin
Simon Burke
2 March 2017 (2017-03-02)
0.180

Khatri pressures Munk into diverting the police's attention to find Anderssen and Elena, despite Tomak just having become the fourth victim of the still unidentified serial killer. Michael tries to convince Natalie and Vincent of the possibilities of full tissue regeneration with the evidence file and films from 1942. Hildur and Eric are forced to confront their feelings for one another, but when Hildur tries to convince Eric of Munk's corruption, he refuses to believe her. Meanwhile, a large sinkhole appears on a highway out on the glacier, and Munk is reluctant to spend the town's resources on fixing it. Petra manages to track down Dan, who then manages to blackmail Khatri to secure his release from custody. Hildur decides to challenge Munk to find out the truth behind her removal as governor, but her efforts to lure him into a trap, resulting in his own downfall, end in tragedy.

19
7
"Episode 7"
Metin Hüseyin
Simon Donald
9 March 2017 (2017-03-09)
0.215

Munk continues to cover up the events surrounding Hildur's death. As Freya's health begins to deteriorate, Michael goes in search of the weather station featured in the video from 1942, but discovers more than he bargained for when he is confronted by a lone Russian soldier. Munk urges Eric to step up the investigation and nail the killer before he takes another victim. Following a heart-to-heart with Vladek, Anderssen crushes Elena to death as he realized he isn't his former self anymore. Natalie approaches Munk with evidence of Khatri's corruption, but he refuses to investigate. Eric's suspicions begin to grow when it becomes apparent that Hildur has been uncontactable for some time. When he receives news of a body found floating in a lake out on the glacier, he is heartbroken to discover that the victim is his estranged wife.

20
8
"Episode 8"
Kieron Hawkes
Simon Burke
16 March 2017 (2017-03-16)
0.174

Two men stumble upon Vladek's shrine cave and are killed. Vincent and Ingrid wake up together. Khatri confronts a frightened Anderssen about his genetic changes, warns him he will experience uncontrollable violence, and offers to cure him; instead he takes over the murder investigations while Eric blames himself for Hildur's "suicide." Michael brings the female Russian soldier's body to Vincent. He tries to convince Eric that Hildur was murdered, but he again refuses to listen. Michael finally brings his suspicions to Ingrid and Petra, who hear him out. Terrified that Natalie knows about the tissue regeneration, Munk and Khatri conspire to steal all her evidence and poison her. Vladek finds Elena's body and burns it Viking style. Anderssen wakes up a lab table but Vladek soon incapacitates her and retrieves the former sheriff.

21
9
"Episode 9"
Kieron Hawkes
Simon Donald
23 March 2017 (2017-03-23)
0.152

Vladek forces Anderssen to take muscimol with him and then reveals the tupilaq. Freya arrives and burns it. Vladek escapes with an unconscious Freya while Michael and Ingrid save Anderssen from the fire. Vincent finds Natalie bleeding from the eyes, apparently infected. Michael and Ingrid discover where and how Hildur was murdered. Vladek castrates himself, cauterizes the wound, and turns himself in to Petra. Anderssen locks Petra in a cell and beats Vladek to death and drags his body into the town hall. Michael takes Munk at gunpoint and Munk promises to save Freya.

22
10
"Episode 10"
Kieron Hawkes
Simon Donald
30 March 2017 (2017-03-30)
0.113

The extraction team arrive in Fortitude in search of Dan. Natalie's condition continues to worsen, but, when Vincent discovers that Khatri is leaving town, he is forced to take drastic action to potentially save his friend's life. The sheriff announces to the townspeople that Hildur was murdered and gives a chilling account of her death, naming Munk as her killer. As he allows the townspeople to carry out their own form of justice, Michael realises that he is the only person who holds the key to saving Freya. As Michael continues his search for answers, Eric comes to his rescue when he is shot by a member of the extraction team.

text: Reception[edit]

text: Critics gave positive reviews. Review aggregator Rotten Tomatoes gave the first season an 89% rating with an average score rating of 7.74 out of 10 based on 27 critic reviews. The critical consensus reads "Fortitude's fine cast, deliberate pacing, and chilling setting provide the show with more than enough of the titular quality."[16] Metacritic gave the first season a score of 75 out of 100, which indicates "generally favorable reviews" based on 15 critic reviews.[17]

text: The second season received generally positive reviews. On Rotten Tomatoes, the season has an 100% approval rating with an average rating of 7.17 out of 10 based on 6 reviews.[18]

text: Broadcast[edit]

text: In the United States and Canada, the series premiered on 29 January 2015 (the same date as the UK) on Pivot and Super Channel, respectively.[19][20] The series debuted in Australia on 15 February 2015 on ABC.[21]

text: Controversy[edit]

text: The violent scenes broadcast of the series by FX on 19 February 2017 in Indonesia led to the banning of the channel by Indonesian authority from 10 to 14 April 2017.[22]


document_title: Hyacinth macaw

question_text: what is the biggest parrot in the world

question_tokens: what is the biggest parrot in the world

text: Hyacinth macaw

text: Taxonomy[edit]

text: English physician, ornithologist, and artist John Latham first described the hyacinth macaw in 1790 under the binomial name Psittacus hyacinthinus based on a taxidermic specimen sent to England.[2] It is one of two extant and one probably extinct species of the South American macaw genus Anodorhynchus.

text: Description[edit]

text: The largest parrot by length in the world, the hyacinth macaw is 100 cm (3.3 ft) long from the tip of its tail to the top of its head and weighs 1.2–1.7 kg (2.6–3.7 lb).[3][4] Each wing is 388–425 mm (15.3–16.7 in) long.[3] The tail is long and pointed.[3] Its feathers are entirely blue, lighter above. However, the neck feathers can sometimes be slightly grey.

text: Behavior[edit]

text: Food and feeding[edit]

text: The majority of the hyacinth macaw diet is nuts, from native palms, such as acuri and bocaiuva palms.[5] They have very strong beaks for eating the kernels of hard nuts and seeds. Their strong beaks are even able to crack coconuts, the large brazil nut pods, and macadamia nuts. The birds also boast dry, smooth tongues with a bone inside them that makes them an effective tool for tapping into fruits.[6] The acuri nut is so hard, the parrots cannot feed on it until it has passed through the digestive system of cattle.[5] In addition, they eat fruits and other vegetable matter. The hyacinth macaw generally eats fruits, nuts, nectar, and various kinds of seeds. Also, they travel for the ripest of foods over a vast area.[7]

text: In the Pantanal, hyacinth macaws feed almost exclusively on the nuts of Acrocomia aculeata and Attalea phalerata palm trees. This behaviour was recorded by the English naturalist Henry Walter Bates in his 1863 book The Naturalist on the River Amazons, where he wrote that

text: It flies in pairs, and feeds on the hard nuts of several palms, but especially of the Mucuja (Acrocomia lasiospatha). These nuts, which are so hard as to be difficult to break with a heavy hammer, are crushed to a pulp by the powerful beak of this macaw.

text: Charles Darwin remarked on Bates's account of the species, calling it a "splendid bird" with its "enormous beak" able to feed on these palm nuts.[9]

text: Tool use[edit]

text: Limited tool use has been observed in both wild and captive hyacinth macaws. Reported sightings of tool use in wild parrots go as far back as 1863. Examples of tool use that have been observed usually involve a chewed leaf or pieces of wood. Macaws often incorporate these items when feeding on harder nuts. Their use allows the nuts the macaws eat to remain in position (prevent slipping) while they gnaw into it. It is not known whether this is learned social behavior or an innate trait, but observation on captive macaws shows that hand-raised macaws exhibit this behavior, as well. Comparisons show that older macaws were able to open seeds more efficiently.[10]

text: Reproduction[edit]

text: Nesting takes place between July and December, with nests constructed in tree cavities or cliff faces depending on the habitat.[11] In the Pantanal region, 90% of nests are constructed in the manduvi tree (Sterculia apetala). The hyacinth macaw depends on its predator, the toucan, for its livelihood. The toucan contributes largely to seed dispersal of the manduvi tree that the macaw needs for reproduction.[12] However, the toucan is responsible for dispersing 83% of the seeds of Sterculia apetala, but also consumes 53% of eggs preyed.[13] Hollows of sufficient size are only found in trees around 60 years of age or older, and competition is fierce.[12] Existing holes are enlarged and then partially filled with wood chips.[14] The clutch size is one or two eggs,[5] although usually only one fledgling survives[5] as the second egg hatches several days after the first, and the smaller fledgling cannot compete with the firstborn for food. A possible explanation for this behavior is what is called the insurance hypothesis. The macaw lays more eggs than can be normally fledged to compensate for earlier eggs that failed to hatch or firstborn chicks that did not survive.[15] The incubation period lasts about a month, and the male tends to his mate whilst she incubates the eggs.[5] The chicks leave the nest, or fledge, around 110 days of age,[citation needed] and remain dependent on their parents until six months of age.[5] They are mature and begin breeding at seven years of age.

text: General traits[edit]

text: Hyacinth macaws are the largest psittacine. They are also very even-tempered and can be calmer than other macaws, being known as "gentle giants".[16] An attending veterinarian must be aware of specific nutritional needs and pharmacologic sensitivities when dealing with them. Possibly due to genetic factors or captive rearing limitations, this species can become neurotic/phobic, which is problematic.[17]

text: Distribution and habitat[edit]

text: The hyacinth macaw survives today in three main populations in South America: In the Pantanal region of Brazil, and adjacent eastern Bolivia and northeastern Paraguay, in the Cerrado region of the eastern interior of Brazil (Maranhão, Piauí, Bahia, Tocantins, Goiás, Mato Grosso, Mato Grosso do Sul, and Minas Gerais), and in the relatively open areas associated with the Tocantins River, Xingu River, Tapajós River, and the Marajó island in the eastern Amazon Basin of Brazil. Smaller, fragmented populations may occur in other areas. It prefers palm swamps, woodlands, and other semiopen, wooded habitats. It usually avoids dense, humid forest, and in regions dominated by such habitats, it is generally restricted to the edge or relatively open sections (e.g. along major rivers). In different areas of their range, these parrots are found in savannah grasslands, in dry thorn forest known as caatinga, and in palm stands,[11] particularly the Moriche Palm (Mauritia flexuosa).[18]

text: Conservation and threats[edit]

text: The hyacinth macaw is an endangered species due to the cage bird trade and habitat loss.[11] In the 1980s, an estimated 10,000 birds were taken from the wild and at least 50% were destined for the Brazilian market.[19] Throughout the macaw's range, habitat is being lost or altered due to the introduction of cattle ranching and mechanised agriculture, and the development of hydroelectric schemes.[11] Annual grass fires set by farmers can destroy nest trees, and regions previously inhabited by this macaw are now unsuitable also due to agriculture and plantations. Locally, it has been hunted for food, and the Kayapo Indians of Gorotire in south-central Brazil use its feathers to make headdresses and other ornaments. While overall greatly reduced in numbers, it remains locally common in the Brazilian Pantanal, where many ranch-owners now protect the macaws on their land.[20]

text: The hyacinth macaw is protected by law in Brazil and Bolivia,[11] and commercial export is banned by its listing on Appendix I of the CITES.[21] A number of long-term studies and conservation initiatives are in place; the Hyacinth Macaw Project in the Brazilian state of Mato Grosso do Sul has carried out important research by ringing individual birds, and has created a number of artificial nests to compensate for the small number of sites available in the region.[5]

text: The Minnesota Zoo with BioBrasil and the World Wildlife Fund are involved in hyacinth macaw conservation.[14][22]

text: Causes of endangerment[edit]

text: Parrots as a whole, being of the family Psittacidae, are some of the most threatened birds in the world. This family has the most endangered species of all bird families, especially in the neotropics, the natural home of the hyacinth macaw, where 46 of 145 species are at a serious risk of global extinction.[23] This species qualifies as endangered on the IUCN Red List because the population has suffered rapid reductions with the remaining threats of illegal trapping for the cage bird trade and habitat loss [24] A few serious threats to the survival of the species in the Pantanal include human activities, mainly those resulting in habitat loss, the burning of land for pasture maintenance, and illegal trapping [25] The exceptionally noisy, fearless, curious, sedentary, and predictable nature of this species, along with its specialization to only one or two species of palm in each part of its range, makes them especially vulnerable to capture, shooting, and habitat destruction.[26] Eggs are also regularly preyed on by corvids, opossums, and coatis.[27] Adults have no known natural predators.[4] The young are parasitized by larvae of flies of the genus Philornis.[28]

text: Although the species has a low genetic variability, it does not necessarily pose a threat to their survival. This genetic structure accentuates the need for protection of hyacinth macaws from different regions to maintain their genetic diversity. Nevertheless, the most important factors negatively affecting the wild population prove to be habitat destruction and nest poaching.[29]

text: In the Pantanal, habitat loss is largely contributed to the creations of pastures for cattle, while in many other regions, it is the result of clearing land for colonization.[30] Similarly, large areas of habitat in Amazonia have been lost for cattle ranching and hydroelectric power schemes on the Tocantins and Xingu Rivers. Many young manduvi trees are then being grazed on by cattle or burnt by fire, and the Gerias is speedily being converted to land for mechanized agriculture, cattle ranching, and exotic tree plantations.[1] Annual grass fires set by farmers destroy a number of nest trees, and the rise of agriculture and plantations has made habitats formerly populated by the macaws unsuitable to maintain their livelihoods.[24] Moreover, increase in commercial demand for feather art by the Kayapo Indians threatens the species, as up to 10 macaws are needed to make a single headdress.[26]

text: In the event of the macaws being taken from their natural environment, a variety of factors alter their health such as inadequate hygiene conditions, feeding, and overpopulation during the illegal practice of pet trade. Once birds are captured and brought into captivity, their mortality rates can become very high.[30] Records reveal a Paraguayan dealer receiving 300 unfeathered young in 1972, with all but three not surviving. Due to the poor survival rates of the young, poachers concentrate more heavily on adult birds, which depletes the population at a rapid pace.[26]

text: According to Article 111 of Bolivian Environmental Law #1333, all persons involved in the trade, capture, and transportation without authorization of wild animals will suffer a two-year prison sentence, along with a fine equivalent to 100% of the value of the animal.[31] While many trackers have been arrested, the illegal pet trade still largely continues in Santa Cruz, Bolivia. Unfortunately, animal trafficking is not necessarily viewed as a priority in the city, leaving national departmental and municipal governments unwilling to halt the trade in city centers, and local police reluctant to get involved. This ideology has in turn resulted in a lack of enforcement regarding trade in both CITES-restricted species and threatened species, with little to no restrictions regarding humane treatment of the animals, disease control, or proper hygiene. In the trade centers, the hyacinth macaw demanded the highest price of US$1,000, proving it to be a very desirable and valued bird in the pet trade industry.[32]

text: Conservation steps taken to preserve the hyacinth macaw[edit]

text: In 1989, the European Endangered Species Programme for the hyacinth macaw was founded as a result of concerns about the status of the wild population and the lack of successful breeding in captivity.[31] Breeding in captivity still remains difficult, being that hand-reared hyacinth macaw offspring have demonstrated to have higher mortality rates, especially within the first month of life. Additionally, they have a higher incidence of acute crop stasis than other macaw species due, in part, to their specific dietary requirements.[33] The hyacinth macaw is protected by law in Brazil and Bolivia, and international trade is prohibited by its listing on Appendix I of the CITES.[24] Appendix I has banned exporting the bird in all countries of origin, and several studies and conservational initiatives have been taken.[1] The Hyacinth Macaw Project in the Caiman Ecological Refuge, located in the Pantanal, has employed artificial nests and chick management techniques, along with effectively raising awareness among cattle ranchers. Many ranch owners in the Pantanal and Gerais, to protect the birds, no longer allow trappers on their properties.[1]

text: A number of conservation actions have been proposed, including the study of the current range, population status, and extent of trading in different parts of its range. Additionally, propositions have been made to assess the effectiveness of artificial nest boxes, enforce legal measures preventing trade, and experiment with ecotourism at one or two sites to encourage donors.[1] Furthermore, the Hyacinth Macaw Project in Mato Grosso do Sul has carried out important research by ringing individual birds and has created a number of artificial nests to compensate for the small number of sites available in the region.[24] Furthermore, proposals to list the species as Endangered under the U.S. Endangered Species Act have been made to further protective measures in the USA and to create Bolivian and Paraguayan trade management authorities under presidential control.[26]

text: Long-term prospects[edit]

text: Each of the three main populations should be managed as a separate biological entity so as to avoid numbers dropping below 500. While the birds may be in decline in the wild, notably higher populations of captive macaws are being held in zoos and private collections. If success in managing and replanting the macaw’s food trees and erecting nest boxes as an experiment in the Pantanals is seen, the species could survive. Survival rates could also be enhanced if ranch owners would leave all large and potential nest trees standing and eliminate all trapping on their properties. Ultimately, should these factors work in tandem with erection of nest boxes, fencing off of certain saplings, and the planting of others, the long-term prospects of the hyacinth macaw species would be greatly improved.[26]

text: Gallery[edit]

text:
Images of hyacinth macaws in the Pantanal, Brazil

Manduvi Cougar Mountain Zoo