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Legion gored Gamera with her sharp legs and blew his shoulder off with her laser beam, then left him for dead as the flower began to seed. Gamera limped to the flower and held himself in front of it when it seeded, preventing the seed from launching but falling victim to a nuclear force explosion that leveled the entire city. Gamera was apparently killed by the explosion, giving the Legion free reign to move on to Tokyo , which was still recovering from the previous year's battle. Several people, including Asagi, surrounded the fallen Gamera, praying for him to recover.

Asagi held out the bead that bonded her to Gamera, which shattered in her hand as huge amounts of energy were absorbed by Gamera's body. Gamera was revived and flew to Tokyo to stop the Legion, while Asagi noticed that her bond with Gamera had been broken. Despite his best efforts, Gamera was unable to defeat Legion. Out of options, Gamera absorbed a large portion of Earth's Mana energy, which gave him enough power to perform his ultimate attack: the Ultimate Mana Blast. Gamera's plastron opened and fired a huge beam of plasma that vaporized the Mother Legion.

Gamera looked at the JSDF forces that had aided him before roaring victoriously and flying away. Realizing Gamera's dedication to defending the Earth from any threat, humanity was left to wonder of the consequences if he ever deemed them a threat to the Earth. Gamera's use of Earth's Mana to revive himself and defeat Legion severely depleted its presence on Earth, allowing clutches of Gyaos eggs across the globe to hatch and evolve into more powerful forms called Hyper Gyaos. Gamera, believing himself responsible for this situation, traveled across the globe aggressively hunting down and eliminating all of the Gyaos he could find.

When two Hyper Gyaos appeared in Shibuya, Gamera arrived and caused horrific destruction and countless deaths in his attempts to kill them. Meanwhile, a girl named Ayana Hirasaka , who blamed Gamera for her parents' deaths during his battle with Super Gyaos in , discovered a strange creature in a shrine, which she named Iris and began to raise in the hopes it would one day kill Gamera and take revenge for her.

When its attempt to bond itself to Ayana failed and she was taken away, Iris rampaged through the village where she lived and killed countless villagers before escaping to the countryside. When Iris detected Ayana was in Kyoto, it began to fly there, attacking J. Gamera arrived and attacked Iris in the air, but was shot out of the sky by the jets, who still believed him the bigger threat.

Iris arrived in Kyoto as a typhoon struck the city, intent on recovering Ayana. When Gamera touched down in the city, Ayana ordered Iris to kill him. Gamera and Iris battled in downtown Kyoto, setting the city ablaze. Iris impaled Gamera with its spear-tipped arm, and the two monsters tackled each other into the Kyoto train station. Gamera fell unconscious from his wounds and appeared dead, leaving Iris to bond with Ayana. Iris forced Ayana into its chest against her will, showing her visions of the people it had killed.

Gamera regained consciousness and interrupted the fusion by punching into Iris' chest and grabbing Ayana. Iris reacted by impaling Gamera's hand against a wall with its spear hand, then drained his blood through it. Iris copied Gamera's plasma fireballs and prepared to fire them at Gamera. Gamera used a fireball to sever his own hand, then caught Iris' fireballs with the stump, using his Mana manipulation powers to form a fiery plasma fist. Gamera plunged this fist into Iris' chest wound, causing the creature to explode from the inside.

Gamera set Ayana down in front of Asagi Kusanagi and her friend Mayumi Nagamine, who tried desperately to revive her. Gamera roared loudly, causing Ayana to awaken. Ayana wondered why Gamera saved her, then began to sob and beg for forgiveness. Sensing a swarm of thousands of Hyper Gyaos approaching, the wounded Gamera exited the station and waited for their arrival. Nagamine remarked that it appeared Gamera would continue to fight, even if alone, while Asagi stated that Gamera wasn't alone, knowing humanity was ready to fight by his side once more.

In , Gamera appeared to save a Japanese village from a swarm of Gyaos. Gamera was severely wounded by the Gyaos' sonic beams and overwhelmed by their numbers, so he self-destructed, taking the Gyaos with him. Gamera's sacrifice was gradually forgotten - The country was now at peace after 33 years without any monsters appearing, and Gamera was seen as a memory of a time when Japan was constantly besieged by disastrous monster attacks.

Over thirty years later, a boy named Toru discovered a turtle egg on a glowing red rock on an island near his home. The egg hatched into a baby turtle that Toru named " Toto ," a nickname his late mother once called him. As Toto grew, he began demonstrating unusual abilities, including flight and the ability to spit fireballs. Toru gradually began to realize that Toto was a baby Gamera, and started to hide him in a barn as he grew increasingly large.

Eventually, Toto escaped, much to Toru's dismay. When a giant sea lizard called Zedus attacked Toru's village, Toto reappeared, now eight meters in height. Toto battled Zedus while Toru and his family escaped, but was brutally maimed and nearly killed by the more powerful Zedus. When Zedus attacked Nagoya , Toto escaped from the facility where he was being held and, now grown to 30 meters in height, attacked Zedus again.

Toto was still no match for Zedus, and was tossed headfirst into a skyscraper. Toru ran to the building and approached Toto, begging for him to not sacrifice himself like his predecessor did years before. Toru presented Toto with the stone his egg had been found on. When Toto ate it, he was re-energized and began to fly, slamming into Zedus and knocking him off the building. Toto and Zedus battled again in the street, with Zedus launching his tongue at Toto. Toto caught the tongue and tore it out, leaving Zedus reeling in pain.

Toto then launched a fireball at Zedus, which collided with the beast's head and killed him in a fiery explosion. As Toto flew off into the horizon, Toru tearfully said "Sayonara In the short film first shown at New York Comic-Con to celebrate Gamera's 50th anniversary, Gamera is shown killing a swarm of Gyaos in Tokyo ten years in the past, at the cost of thousands of human lives.

In the present day, a new tentacled kaiju appears in Tokyo, and Gamera arrives to battle it. Gamera's shell is extremely resilient and strong. Missiles and other weaponry merely bounce off of it, along with most of his opponents' attacks. There have been a few times where his shell has faltered, most notably when Guiron hammered at the same spot several times and began cutting through.

Gyaos ' sonic Beam, Zigra 's Paralyzing Beam, and Barugon 's rainbow ray cannot penetrate Gamera's shell, shown in the films when he withdraws into his shell to avoid the attacks. This is most visible when Viras impales his stomach with the tip of its head, and when Iris impales Gamera straight through his entire body by stabbing him through the stomach. In the original film , it was said that Gamera's cellular tissue was so dense that conventional weapons were useless against him. However, in the Heisei series, tanks were able to injure Gamera to some extent, while he was easily knocked out of the sky by missiles.

In the Showa series, Gamera fed on fire and was attracted by other heat sources, such as power plants and Barugon 's "rainbow" ray attack, and in his first movie he was lured to a rocket by fire. He could breathe intensely hot streams of flames from his mouth when caught in a dangerous situation. The Heisei version, on the other hand, could fire explosive plasma fireballs from his mouth, usually very quickly, and with varying accuracy.

At the end of Gamera: Guardian of the Universe , Gamera absorbs fire and fires a "Super-powered" plasma fireball and in Gamera 2: Attack of the Legion , he breathes in the oxygen produced by a Legion plant and fires an "Oxygen-Powered" plasma fireball. The Heisei version could also absorb and manipulate "Mana", or the living essence and energy of life on Earth, and release an extremely powerful stream of pure plasma and fire from an opened, organic "cannon" in his chest.

In Gamera 3: Revenge of Iris , Gamera blasted his own arm off and absorbed plasma fireballs shot by Iris and used his stump arm to grow back his arm in a plasma form. From this, it appears Gamera has the ability to manipulate fire. However, as shown in this film, this ability to regenerate limbs in plasma form seems to be only temporary, as the plasma arm disappeared after defeating Iris.

Perhaps most famously, Gamera also has the ability to fly. Generally Gamera will pull his arms, legs, head, and tail into his shell, fire flames out of his arm and leg cavities and spin around like a frisbee. This mode of flight had an added advantage in the later films, where he would use the sharp edges of his shell to cut enemies while spinning, similar to a circular saw. In the Heisei era films, Gamera's arms would extend and stretch out into wings similar to the flippers of a sea turtle while using this form of flight, giving him added aerodynamics and control.

The Heisei films gave Gamera one more additional weapon: a pair of sharp spikes protruding from his elbows. In his first Heisei era appearance, these spikes were hidden during the majority of the film, extending only when needed in battle. In later appearances they were permanently extended. When seriously or gravely injured, Gamera can enter a coma-like state in order to heal. This often fools his opponents into thinking that he is dead. This ability has been used in almost every film. Gamera's only major weakness is cold. The monster Barugon was able to achieve success against Gamera using his freezing spray, and scientists nearly defeated Gamera during his first appearance using special cadmium freezing bombs.

This weakness was only shown in Gamera's earliest films, and has not been explored since. Gamera's internal organs are also vulnerable to parasitic actions, as shown when he fainted and turned a deathly white when the baby Jiger was growing in his lungs. Gamera appears on the game's second stage, the second part of the tenth stage, and sixteenth stage. On the second and sixteenth stages, Gamera deals with a swarm of Soldier Legion and Legion Plant, while on the second part of the the tenth stage, he exterminates young Gyaos.

Gamera in City Shrouded in Shadow. One year after defeating Super Gyaos , Gamera was sleeping in the ocean as a shark swam toward his closed eye. However, sensing a Gyaos attack, Gamera awoke, scaring away the shark, and flew up from the ocean to Guanajota, Mexico. Upon locating Gyaos, Gamera flew up behind the beast and blasted it with a fireball.

The creature turned around to face its aggressor and returned fire. Gyaos then sprang on Gamera, and knocked him to the ground. Gamera however, was able to use one of his tusks to stab Gyaos' shin, which caused it to retreat long enough for Gamera to stand back up as the frightened people fled the ruined city. Gyaos charged at Gamera with a headbutt, but Gamera grabbed Gyaos' shoulders and lead it into a roll that Gamera used to slam Gyaos into the water.

Gyaos managed to get back up, but it then saw Gamera blasting through the air toward it. Gamera tackled Gyaos so that it fell backwards into a tanker ship in the harbor, and then blasted upward and spat his fire blast at Gyaos, igniting the oil in the tanker and bringing the entire Guanajota harbor to a boil. Gamera sustained the blast on the harbor, and the helpless Gyaos clone was effectively obliterated. With his job done, Gamera then turned around and began flying out over the Pacific Ocean. While flying, Gamera became aware of the alien Zigra attacking an envoy of French warships, and took action against it.

Gamera blasted Zigra with a fireball, and began to speed toward it for an attack, but Zigra hit Gamera with its Tri-Color Ray Blast, and Gamera was incapacitated and began to sink into the ocean. Before Gamera hit the bottom, a French submarine fired at his head, causing him to wake up. Gamera burst through the surface of the water faster than Zigra could react, and bit the beast in its throat. Gamera then flew away with the helpless Zigra in his jaws. Later, in Paris , France , Gamera flew in under the command of Dr.

Greta Karbone , who held the Atlantean Magatama , and through Viras' command could use it to control Gamera. Gamera then began to fly into planes and missiles and other types of fire in order to defend Viras during its destruction of the city. Gamera continued to defend Viras, and flew headfirst into a fighter jet, the impact from which caused Gamera to bleed from his mouth. Gamera willingly threw himself into the onslaught of incoming fire until Karbone was separated from the Magatama.

Then, with his shell smoking from his defenselessly having sustained endless attacks, he turned around in the air and prepared to blast fire at Viras.

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However, Viras shot its electric ray at him, and the weakened Gamera went down. He began to try to crawl toward his adversary, but was too drained to continue.

However a man by the name of Gusano , out of love for the possessed Karbone, tore out an alien spaceship thruster that was about to explode, and threw it at Gamera, warning him that the fire he needed to survive was as dangerous and all-consuming as Gusano's love for Karbone had been. Gusano died in the explosion, but it proved to be just what Gamera needed.

Reinvigorated, Gamera picked up a building and threw it at Viras, despite its attacks. Gamera then bit one of Viras' tentacles and with one flip of his head, sent the beast flying. It landed on one of the steeples of the Cathedral of Notre Dame. With the day won, Gamera flew away. Gamera appears in the Gamera vs.

The comic serves as a retelling of Gamera vs. Barugon to incorporate it into the Heisei timeline. In the spin-off manga to Gamera the Brave , Gamera's battle against the Gyaos in is gone more into detail within the "Special Link" chapter. Gamera is being attacked by 4 Gyaos' laser beams. Gamera kills a first Gyaos with a point-blank fire blast and then stomps on its head. The 3 Gyaos keep firing at Gamera, but Gamera gets in his shell and charges up a fire blast which he fires at one of the Gyaos.

Another Gyaos comes in and bites Gamera's arm, and Gamera throws it to the ground and fires at it, killing it. The 2 remaining Gyaos hover about, and Gamera fires another fire blast at one of them. From the smoke the Gyaos flies toward Gamera and unleashes a flurry of beams. Gamera punches it to the ground. However, the Gyaos continues its onslaught of beams just as the other Gyaos turned out to have survived and flies over to Gamera.

Gamera begins spurting blood , and the two Gyaos feast on Gamera's entrails. Gamera, acknowledging he's been defeated, triggers his Fireball Ejection Suicide, killing both him and both Gyaos. Some of Gamera's roars would later be reused for the Godzillasaurus. This is a list of references for Gamera. These citations are used to identify the reliable sources on which this article is based. These references appear inside articles in the form of superscript numbers, which look like this: [1].

Showing 66 comments. When commenting, please remain respectful of other users, stay on topic, and avoid role-playing and excessive punctuation. Comments which violate these guidelines may be removed by administrators. Gamera is really neat! Gamera is filled with meat! We've been eating Ga-mer-aaaaaaa!!! Astounding Beyond Belief. Titan of Water. Human: cause you destroy cities when your hungry Gamera eats plasma filled Snickers Human: Friend of all Children? Gorosaurus Fan. Keizer Zilla. The King of the Monsters. Chainsaw Gigan.

Green Blob Thing. Emperor Ghidorah. Shin Varan. Welcome to Wikizilla! Log in or sign up to edit pages and contribute. Toto Showa Heisei Avant. We commit to the cradle of time the Last Hope, Gamera. May he awaken with the Shadow of Evil, Gyaos. Gamera in Gamera: Battle. For additonal information or feedback, visit help. You 0. Collaboration with Kaiju! This is the big batch version of a one off keg… Read More.

This is the big batch version of a one off keg brewed at earlier in Packaged in keg and ml bottles on 20 August Read Less. Schwarzbier "shvahrts-beer" , is simply German for black beer. Imperial is just the way we all like our black German lagers. Inappropriate Touching is… Read More. Inappropriate Touching is just our way to remember what was going on during the making of this beer.

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Read Less. In the deep southeast a butterfly flaps its wings; two weeks later in the subtropical north a tornado is born. Such is the chaos that surrounds this Double… Read More. Such is the chaos that surrounds this Double India Pale Ale. A chance meeting, a quick friendship and two regional breweries, coming together to celebrate the sensitive dependence on initial conditions in a deterministic nonlinear system. Also, beer was involved. A decent cocoon of malted wheat has dried out the body and allowed the hops to break free. Getting jammed in all over the place, we have Simcoe, Citra, Cascade, Mosaic and Enigma, all playing their parts in a tropical, piney, dank, resinous, lupulin bomb.

Have a sip, have two, who knows where this will lead. Have you ever seen a portal? This is what cames out of two brewers… Read More. This is what cames out of two brewers trying to one up each other when designing a recipe. This beer might just have had too many hops and too much malt, especially rye. Not being able to leave well enough alone they brewed it again for Good Beer Week, this time it is for the rest of Australia. Four students, four breweries, four hops, four timepoints, it's a 4 Way IPA.

With Croft, Dainton, Kaiju and Newstead all meeting at beer school it was only a matter of time until the alumni got together to make a ridiculous beer.

Fast and sensitive taxonomic classification for metagenomics with Kaiju

Using Azacca at 15 minutes, Riwaka at 5 minutes, Motueka in the Whirlpool and Columbus as a double dry-hop this monster is all about excess. Could it be Dr Aldred's biggest mistake? Almost definitely, and we couldn't be happier. Taking none of the lessons we learnt about balance and control we pushed things to the edge to create what we are calling a quadruple IPA.

Once a match is found, which has a higher score than all remaining fragments in the search list and a score above the minimum score threshold s , the search stops and this highest scoring match is used for classifying the read.

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If multiple matches to several different database entries have the same score, Kaiju classifies the read to their LCA as above. Kaiju outputs one line for each read or read pair , containing the read name and the NCBI taxon identifier of the assigned taxon, as well as the length or score of the match. Optionally, Kaiju can also produce a summary file with the number of reads assigned per taxon, which can be loaded into Krona 26 for interactive visualization.

We also include a utility program that can merge the classification results from different runs or programs, for example, for merging Kaiju and Kraken results. The primary goal of Kaiju's protein-level classification is to improve classification of those parts of a metagenome that are only distantly related to the known sequences or belong to a branch of the phylogeny that is underrepresented in the reference database. We therefore devised a benchmark study, which addresses this problem by simulating the classification of metagenomic reads from a novel strain or species that is not contained in the reference database.

Only those genomes were retained that are assigned to a species belonging to a genus and have a full chromosome with annotated proteins, resulting in a total of 2, genomes belonging to distinct genera. For example, the genus Streptococcus contains genomes, whereas genera have only 1 available genome, genera have 2 available genomes and so on. The distribution clearly illustrates a sampling bias and the sparseness across large parts of the phylogeny. From the total of 2, genomes, we extracted those genera that have at least 2 and at most 10 genomes assigned.

This resulted in a list of genera comprising genomes, for which we measured the classification performance individually. For each of the genomes, we simulated five sets of HTS reads and created a reference database not containing this genome that is then used to classify the simulated reads. Reads were simulated from the whole genome including plasmids using ART The four sets of Illumina reads contain 50k reads of length either or nt, both in single-end and paired-end mode.

To evaluate classification accuracy, we measured the number of classified reads, as well as sensitivity and precision on genus- and phylum-levels. The same measurements were used in the study by Ounit et al.

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Kraken v0. Performance was measured in processed reads or read pairs per second r. While Kaiju and Clark need to preload their index into memory before the classification starts, Kraken can either preload the index or only load necessary segments during the classification. We therefore measured Kraken's speed using both options, and it turned out that Kraken runs faster without preloading on our hardware. We therefore report its performance without preloading. For each of the five types of simulated reads from our exclusion benchmark, we created a data set comprising 10k reads from each genome in the reference database, resulting in Figures were made with the ggplot2 package 28 in the R statistical software How to cite this article: Menzel, P.

Fast and sensitive taxonomic classification for metagenomics with Kaiju. Author contributions P. National Center for Biotechnology Information , U. Nat Commun. Published online Apr Author information Article notes Copyright and License information Disclaimer. Received Nov 26; Accepted Mar 7. All Rights Reserved. This work is licensed under a Creative Commons Attribution 4. The images or other third party material in this article are included in the article's Creative Commons license, unless indicated otherwise in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material.

This article has been cited by other articles in PMC. Abstract Metagenomics emerged as an important field of research not only in microbial ecology but also for human health and disease, and metagenomic studies are performed on increasingly larger scales. Results Protein-level sequence classification Kaiju translates metagenomic sequencing reads into the six possible reading frames and searches for maximum exact matches MEMs of amino acid sequences in a given database of annotated proteins from microbial reference genomes. Genome exclusion benchmark Benchmarking a classifier's accuracy can be done by simulation studies, which, knowing the ground truth about the origin of the simulated reads, can assess the sensitivity and precision of the classification.

Open in a separate window. Figure 1. Genus-level sensitivity and precision. Figure 2. Average sensitivity and precision. Real metagenomes To assess how many reads can actually be classified in real metagenomic data sets, we arbitrarily selected 10 previously published data sets from different microbiomes that were sequenced using various different HTS instruments. Figure 3. Classification of real metagenomes. HiSeq and MiSeq mock communities In addition to the real metagenomes, we also measured Kaiju's and Kraken's performance using the same metrics and reference database on the HiSeq and MiSeq mock community data sets from previous benchmarks 8 , 9.

Runtime and memory The read data set for the runtime benchmark contained Figure 4. Classification speed. Discussion When performing sequence comparison, as in the case of taxonomic assignment using a reference database, there is the obvious trade-off between an algorithm's speed and accuracy. Methods Metagenome classifier Kaiju classifies individual metagenomic reads using a reference database comprising the annotated protein-coding genes of a set of microbial genomes.

Figure 5. Kaiju's algorithm.

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Performance evaluation The primary goal of Kaiju's protein-level classification is to improve classification of those parts of a metagenome that are only distantly related to the known sequences or belong to a branch of the phylogeny that is underrepresented in the reference database.

Additional information How to cite this article: Menzel, P. Footnotes Author contributions P. References Riesenfeld C. Metagenomics: genomic analysis of microbial communities. Next-generation sequencing technologies for environmental DNA research. Computational meta'omics for microbial community studies. The human gut microbiome: implications for future health care.

The oral microbiome in health and disease.

Tools for mapping high-throughput sequencing data. Bioinformatics 28 , — Scalable metagenomic taxonomy classification using a reference genome database. Bioinformatics 29 , — Kraken: ultrafast metagenomic sequence classification using exact alignments. Genome Biol. CLARK: fast and accurate classification of metagenomic and genomic sequences using discriminative k-mers. BMC Genomics 16 , Detection of low-abundance bacterial strains in metagenomic data sets by eigengenome partitioning.

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Comparative metagenomics of eight geographically remote terrestrial hot springs. Structure and function of the global ocean microbiome. Science , Comparative genomic structure of prokaryotes. Basic local alignment search tool. RAPSearch2: a fast and memory-efficient protein similarity search tool for next-generation sequencing data. Methods 12 , 59—60 An evaluation of the accuracy and speed of metagenome analysis tools.

Fast and accurate short read alignment with Burrows—Wheeler transform. Bioinformatics 25 , — Brahme A. Liu Y. Long read alignment based on maximal exact match seeds. Bioinformatics 28 , i—i Jiang M. BMC Bioinformatics 9 , Interactive metagenomic visualization in a web browser.