francesco redi experiment summary

3.1 Spontaneous Generation

Learning objectives.

By the end of this section, you will be able to:

• Explain the theory of spontaneous generation and why people once accepted it as an explanation for the existence of certain types of organisms
• Explain how certain individuals (van Helmont, Redi, Needham, Spallanzani, and Pasteur) tried to prove or disprove spontaneous generation

Clinical Focus

Barbara is a 19-year-old college student living in the dormitory. In January, she came down with a sore throat, headache, mild fever, chills, and a violent but unproductive (i.e., no mucus) cough. To treat these symptoms, Barbara began taking an over-the-counter cold medication, which did not seem to work. In fact, over the next few days, while some of Barbara’s symptoms began to resolve, her cough and fever persisted, and she felt very tired and weak.

• What types of respiratory disease may be responsible?

Jump to the next Clinical Focus box

Humans have been asking for millennia: Where does new life come from? Religion, philosophy, and science have all wrestled with this question. One of the oldest explanations was the theory of spontaneous generation, which can be traced back to the ancient Greeks and was widely accepted through the Middle Ages.

The Theory of Spontaneous Generation

The Greek philosopher Aristotle (384–322 BC) was one of the earliest recorded scholars to articulate the theory of spontaneous generation , the notion that life can arise from nonliving matter. Aristotle proposed that life arose from nonliving material if the material contained pneuma (“spirit” or “breath”). As evidence, he noted several instances of the appearance of animals from environments previously devoid of such animals, such as the seemingly sudden appearance of fish in a new puddle of water. 1

This theory persisted into the 17th century, when scientists undertook additional experimentation to support or disprove it. By this time, the proponents of the theory cited how frogs simply seem to appear along the muddy banks of the Nile River in Egypt during the annual flooding. Others observed that mice simply appeared among grain stored in barns with thatched roofs. When the roof leaked and the grain molded, mice appeared. Jan Baptista van Helmont , a 17th century Flemish scientist, proposed that mice could arise from rags and wheat kernels left in an open container for 3 weeks. In reality, such habitats provided ideal food sources and shelter for mouse populations to flourish.

However, one of van Helmont’s contemporaries, Italian physician Francesco Redi (1626–1697), performed an experiment in 1668 that was one of the first to refute the idea that maggots (the larvae of flies) spontaneously generate on meat left out in the open air. He predicted that preventing flies from having direct contact with the meat would also prevent the appearance of maggots. Redi left meat in each of six containers ( Figure 3.2 ). Two were open to the air, two were covered with gauze, and two were tightly sealed. His hypothesis was supported when maggots developed in the uncovered jars, but no maggots appeared in either the gauze-covered or the tightly sealed jars. He concluded that maggots could only form when flies were allowed to lay eggs in the meat, and that the maggots were the offspring of flies, not the product of spontaneous generation.

In 1745, John Needham (1713–1781) published a report of his own experiments, in which he briefly boiled broth infused with plant or animal matter, hoping to kill all preexisting microbes. 2 He then sealed the flasks. After a few days, Needham observed that the broth had become cloudy and a single drop contained numerous microscopic creatures. He argued that the new microbes must have arisen spontaneously. In reality, however, he likely did not boil the broth enough to kill all preexisting microbes.

Lazzaro Spallanzani (1729–1799) did not agree with Needham’s conclusions, however, and performed hundreds of carefully executed experiments using heated broth. 3 As in Needham’s experiment, broth in sealed jars and unsealed jars was infused with plant and animal matter. Spallanzani’s results contradicted the findings of Needham: Heated but sealed flasks remained clear, without any signs of spontaneous growth, unless the flasks were subsequently opened to the air. This suggested that microbes were introduced into these flasks from the air. In response to Spallanzani’s findings, Needham argued that life originates from a “life force” that was destroyed during Spallanzani’s extended boiling. Any subsequent sealing of the flasks then prevented new life force from entering and causing spontaneous generation ( Figure 3.3 ).

Check Your Understanding

• Describe the theory of spontaneous generation and some of the arguments used to support it.
• Explain how the experiments of Redi and Spallanzani challenged the theory of spontaneous generation.

Disproving Spontaneous Generation

The debate over spontaneous generation continued well into the 19th century, with scientists serving as proponents of both sides. To settle the debate, the Paris Academy of Sciences offered a prize for resolution of the problem. Louis Pasteur , a prominent French chemist who had been studying microbial fermentation and the causes of wine spoilage, accepted the challenge. In 1858, Pasteur filtered air through a gun-cotton filter and, upon microscopic examination of the cotton, found it full of microorganisms, suggesting that the exposure of a broth to air was not introducing a “life force” to the broth but rather airborne microorganisms.

Later, Pasteur made a series of flasks with long, twisted necks (“swan-neck” flasks), in which he boiled broth to sterilize it ( Figure 3.4 ). His design allowed air inside the flasks to be exchanged with air from the outside, but prevented the introduction of any airborne microorganisms, which would get caught in the twists and bends of the flasks’ necks. If a life force besides the airborne microorganisms were responsible for microbial growth within the sterilized flasks, it would have access to the broth, whereas the microorganisms would not. He correctly predicted that sterilized broth in his swan-neck flasks would remain sterile as long as the swan necks remained intact. However, should the necks be broken, microorganisms would be introduced, contaminating the flasks and allowing microbial growth within the broth.

Pasteur’s set of experiments irrefutably disproved the theory of spontaneous generation and earned him the prestigious Alhumbert Prize from the Paris Academy of Sciences in 1862. In a subsequent lecture in 1864, Pasteur articulated “ Omne vivum ex vivo ” (“Life only comes from life”). In this lecture, Pasteur recounted his famous swan-neck flask experiment, stating that “…life is a germ and a germ is life. Never will the doctrine of spontaneous generation recover from the mortal blow of this simple experiment.” 4 To Pasteur’s credit, it never has.

• How did Pasteur’s experimental design allow air, but not microbes, to enter, and why was this important?
• What was the control group in Pasteur’s experiment and what did it show?
• 1 K. Zwier. “Aristotle on Spontaneous Generation.” http://www.sju.edu/int/academics/cas/resources/gppc/pdf/Karen%20R.%20Zwier.pdf
• 2 E. Capanna. “Lazzaro Spallanzani: At the Roots of Modern Biology.” Journal of Experimental Zoology 285 no. 3 (1999):178–196.
• 3 R. Mancini, M. Nigro, G. Ippolito. “Lazzaro Spallanzani and His Refutation of the Theory of Spontaneous Generation.” Le Infezioni in Medicina 15 no. 3 (2007):199–206.
• 4 R. Vallery-Radot. The Life of Pasteur , trans. R.L. Devonshire. New York: McClure, Phillips and Co, 1902, 1:142.

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Lessons and Courses on Microbiology

FRANCESCO REDI (1626-1697)

Francesco Redi, an Italian scientist was the first scientist to challenge the theory of spontaneous generation by demonstrating that living organisms did not actually originate from non-living things. He developed a scientific experiment to test the spontaneous creation of maggots from fresh meats using two jars (one of the jars was left open while the other was closed).

Redi was famously known for his work on spontaneous generation or abiogenesis . He challenged the concept of abiogenesis by showing that maggots on decaying meat came from fly eggs deposited on the meat and not from the meat itself. Redi explained that flies land on exposed meat and lay their eggs which eventually hatch to produce maggots.

Redi performed series of experiments in the early 1670’s in which he covered jars of meat with fine lace that prevented the entry of flies into the jars. Because the meat was covered, no maggots were produced, and this led Francesco Redi to drop the notion of spontaneous generation.

Francesco Redisuccessfully challenged and refuted the theory of spontaneous generation through his work on maggot and flies, in which he showed that maggots on meat came from egg flies. Though his work was known, the ideaof spontaneous generation was not dropped as other scientist like John Needham continued from where he stopped to unravel the mystery behind it.

Barrett   J.T (1998).  Microbiology and Immunology Concepts.  Philadelphia,   PA:  Lippincott-Raven Publishers. USA.

Beck R.W (2000). A chronology of microbiology in historical context. Washington, D.C.: ASM Press.

Brooks G.F., Butel J.S and Morse S.A (2004). Medical Microbiology, 23 rd edition. McGraw Hill Publishers. USA. Pp. 248-260.

Chung K.T, Stevens Jr., S.E and Ferris D.H (1995). A chronology of events and pioneers of microbiology. SIM News , 45(1):3–13.

Slonczewski J.L, Foster J.W and Gillen K.M (2011). Microbiology: An Evolving Science. Second edition. W.W. Norton and Company, Inc, New York, USA.

Summers W.C (2000). History of microbiology. In Encyclopedia of microbiology, vol. 2, J. Lederberg, editor, 677–97. San Diego: Academic Press.

Talaro, Kathleen P (2005). Foundations in Microbiology. 5 th edition. McGraw-Hill Companies Inc., New York, USA.

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2.1 Spontaneous Generation

Learning objectives.

• Explain the theory of spontaneous generation and why people once accepted it as an explanation for the existence of certain types of organisms
• Explain how certain individuals (van Helmont, Redi, Needham, Spallanzani, and Pasteur) tried to prove or disprove spontaneous generation

Humans have been asking for millennia: Where does new life come from? Religion, philosophy, and science have all wrestled with this question. One of the oldest explanations was the theory of spontaneous generation, which can be traced back to the ancient Greeks and was widely accepted through the Middle Ages.

The Theory of Spontaneous Generation

The Greek philosopher Aristotle (384–322 BC) was one of the earliest recorded scholars to articulate the theory of spontaneous generation , the notion that life can arise from nonliving matter. Aristotle proposed that life arose from nonliving material if the material contained pneuma (“vital heat”). As evidence, he noted several instances of the appearance of animals from environments previously devoid of such animals, such as the seemingly sudden appearance of fish in a new puddle of water. [1]

This theory persisted into the 17th century, when scientists undertook additional experimentation to support or disprove it. By this time, the proponents of the theory cited how frogs simply seem to appear along the muddy banks of the Nile River in Egypt during the annual flooding. Others observed that mice simply appeared among grain stored in barns with thatched roofs. When the roof leaked and the grain molded, mice appeared. Jan Baptista van Helmont, a 17th century Flemish scientist, proposed that mice could arise from rags and wheat kernels left in an open container for 3 weeks. In reality, such habitats provided ideal food sources and shelter for mouse populations to flourish.

However, one of van Helmont’s contemporaries, Italian physician Francesco Redi (1626–1697), performed an experiment in 1668 that was one of the first to refute the idea that maggots (the larvae of flies) spontaneously generate on meat left out in the open air. He predicted that preventing flies from having direct contact with the meat would also prevent the appearance of maggots. Redi left meat in each of six containers ( Figure 2 .2 ). Two were open to the air, two were covered with gauze, and two were tightly sealed. His hypothesis was supported when maggots developed in the uncovered jars, but no maggots appeared in either the gauze-covered or the tightly sealed jars. He concluded that maggots could only form when flies were allowed to lay eggs in the meat, and that the maggots were the offspring of flies, not the product of spontaneous generation.

In 1745, John Needham (1713–1781) published a report of his own experiments, in which he briefly boiled broth infused with plant or animal matter, hoping to kill all preexisting microbes. [2] He then sealed the flasks. After a few days, Needham observed that the broth had become cloudy and a single drop contained numerous microscopic creatures. He argued that the new microbes must have arisen spontaneously. In reality, however, he likely did not boil the broth enough to kill all preexisting microbes.

Lazzaro Spallanzani (1729–1799) did not agree with Needham’s conclusions, however, and performed hundreds of carefully executed experiments using heated broth. [3] As in Needham’s experiment, broth in sealed jars and unsealed jars was infused with plant and animal matter. Spallanzani’s results contradicted the findings of Needham: Heated but sealed flasks remained clear, without any signs of spontaneous growth, unless the flasks were subsequently opened to the air. This suggested that microbes were introduced into these flasks from the air. In response to Spallanzani’s findings, Needham argued that life originates from a “life force” that was destroyed during Spallanzani’s extended boiling. Any subsequent sealing of the flasks then prevented new life force from entering and causing spontaneous generation ( Figure 2 .3 ).

• Describe the theory of spontaneous generation and some of the arguments used to support it.
• Explain how the experiments of Redi and Spallanzani challenged the theory of spontaneous generation.

Disproving Spontaneous Generation

The debate over spontaneous generation continued well into the 19th century, with scientists serving as proponents of both sides. To settle the debate, the Paris Academy of Sciences offered a prize for resolution of the problem. Louis Pasteur, a prominent French chemist who had been studying microbial fermentation and the causes of wine spoilage, accepted the challenge. In 1858, Pasteur filtered air through a gun-cotton filter and, upon microscopic examination of the cotton, found it full of microorganisms, suggesting that the exposure of a broth to air was not introducing a “life force” to the broth but rather airborne microorganisms.

Later, Pasteur made a series of flasks with long, twisted necks (“swan-neck” flasks), in which he boiled broth to sterilize it ( Figure 2 .4 ). His design allowed air inside the flasks to be exchanged with air from the outside, but prevented the introduction of any airborne microorganisms, which would get caught in the twists and bends of the flasks’ necks. If a life force besides the airborne microorganisms were responsible for microbial growth within the sterilized flasks, it would have access to the broth, whereas the microorganisms would not. He correctly predicted that sterilized broth in his swan-neck flasks would remain sterile as long as the swan necks remained intact. However, should the necks be broken, microorganisms would be introduced, contaminating the flasks and allowing microbial growth within the broth.

Pasteur’s set of experiments irrefutably disproved the theory of spontaneous generation and earned him the prestigious Alhumbert Prize from the Paris Academy of Sciences in 1862. In a subsequent lecture in 1864, Pasteur articulated “ Omne vivum ex vivo ” (“Life only comes from life”). In this lecture, Pasteur recounted his famous swan- neck flask experiment, stating that “…life is a germ and a germ is life. Never will the doctrine of spontaneous generation recover from the mortal blow of this simple experiment.” [4] To Pasteur’s credit, it never has.

• How did Pasteur’s experimental design allow air, but not microbes, to enter, and why was this important?
• What was the control group in Pasteur’s experiment and what did it show?
• K. Zwier. “Aristotle on Spontaneous Generation.” http://www.sju.edu/int/academics/cas/resources/gppc/pdf/Karen%20R.%20Zwier.pdf ↵
• E. Capanna. “Lazzaro Spallanzani: At the Roots of Modern Biology.” Journal of Experimental Zoology 285 no. 3 (1999):178–196. ↵
• R. Mancini, M. Nigro, G. Ippolito. “Lazzaro Spallanzani and His Refutation of the Theory of Spontaneous Generation.” Le Infezioni in Medicina 15 no. 3 (2007):199–206. ↵
• R. Vallery-Radot. The Life of Pasteur, trans. R.L. Devonshire. New York: McClure, Phillips and Co, 1902, 1:142. ↵

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1-6 spontaneous generation was an attractive theory to many people, but was ultimately disproven..

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Learning Objectives

After reading this section, students will be able to...

• Explain why people believed in the concept of spontaneous generation, the creation of life from organic matter.
• Describe the experiment by Francesco Redi disproved spontaneous generation that disproved spontaneous generation for macroorganisms.
• Explain how did John Needham's experiment re-ignited the debate about spontaneous generation for microorganisms.
• Describe the swan-neck flask experiment of Louis Pasteur and why this ended the debate about spontaneous generation.

Spontaneous generation hypothesizes that some vital force contained in or given to organic matter can create living organisms from inanimate objects. Spontaneous generation was a widely held belief throughout the middle ages and into the latter half of the 19 th century. Some people still believe in it today. The idea was attractive because it meshed nicely with the prevailing religious views of how God created the universe. There was a strong bias to legitimize the idea because this vital force was considered a strong proof of God's presence in the world. Proponents offered many recipes and experiments in proof. To create mice, mix dirty underwear and wheat grain in a bucket and leave it open outside. In 21 days or less, you would have mice. The real cause may seem obvious from a modern perspective, but to the supporters of this idea, the mice spontaneously arose from the wheat kernels.

Another often-used example was the generation of maggots from meat left in the open. Francesco Redi revealed the failing here in 1668 with a classic experiment. Redi suspected that flies landing on the meat laid eggs that eventually grew into maggots . To test this idea, he devised the experiment shown in Figure 1.11. Here he used three pieces of meat. Redi placed one piece of meat under a piece of paper. The flies could not lay eggs onto the meat, and no maggots developed. The second piece was left in the open air, resulting in maggots. In the final test, Redi overlayed the third piece of meat with cheesecloth. The flies could lay the eggs into the cheesecloth, and when he removed this, no maggots developed. However, if Redi placed the cheesecloth containing the eggs on a fresh piece of meat, maggots developed, showing it was the eggs that "caused" maggots and not spontaneous generation. Redi ended the debate about spontaneous generation for large organisms. However, spontaneous generation was so seductive a concept that even Redi believed it was possible in other circumstances.

Figure 1.11. The Redi experiment. . Using several pieces of meat, paper and cheesecloth, Francesco Redi produced compelling evidence against the theory of spontaneous generation. One of the strong points of this experiment was its simplicity, which allowed others to easily reproduce it for themselves. See the text for details of the experiment.

The concept and the debate were revived in 1745 by the experiments of John Needham. It was known at the time that heat was lethal to living organisms. Needham theorized that if he took chicken broth and heated it, all living things in it would die. After heating some broth, he let a flask cool and sit at a constant temperature. The development of a thick turbid solution of microorganisms in the flask was strong proof to Needham of the existence of spontaneous generation. Lazzaro Spallanzani later repeated the experiments of Needham, but removed air from the flask, suspecting that the air was providing a source of contamination. No growth occurred in Spallanzani's flasks, and he took this as evidence that Needham was wrong. Proponents of spontaneous generation discounted the experiment by asserting that the vital force needed air to work properly.

It was not until almost 100 years later that the great French chemist Louis Pasteur, pictured in Figure 1.12, put the debate to rest. He first showed that the air is full of microorganisms by passing air through gun cotton filters. The filter trapped tiny particles floating in the air. By dissolving the cotton with an ether/alcohol mixture, the particles were released and then settled to the bottom of the liquid. Inspection of this material revealed numerous microbes that resembled the types of bacteria often found in putrefying media. Pasteur realized that if these bacteria were present in the air, they would likely land on and contaminate any exposed material.

Figure 1.12. Louis Pasteur . The French microbiologist Louis Pasteur. Drawing by Tammi Henke

Pasteur then entered a contest sponsored by The French Academy of Sciences to disprove the theory of spontaneous generation. Similar to Spallanzani's experiments, Pasteur's experiment, pictured in Figure 1.13, used heat to kill the microbes but left the end of the flask open to the air. In a simple but brilliant modification, he heated the neck of the flask to melting and drew it out into a long S-shaped curve, preventing the dust particles and their load of microbes from ever reaching the flask. After prolonged incubation, the flasks remained free of life and ended the debate for most scientists.

Figure 1.13. The swan neck flask experiment . Pasteur filled a flask with medium, heated it to kill all life, and then drew out the neck of the flask into a long S shape. This prevented microorganisms in the air from easily entering the flask, yet allowed some air interchange. If the swan neck was broken, microbes readily entered the flask and grew

A final footnote on the topic was added when John Tyndall show ed the existence of heat-resistant spores in many materials. Boiling does not kill these spores, and their presence in chicken broth, as well as many other materials, explains the results of Needham's experiments.

While this debate may seem silly from a modern perspective, remember that the scientists of the time had little knowledge of microorganisms. Koch would not isolate microbes until 1881. The proponents of spontaneous generation were neither sloppy experimenters nor stupid. They did careful experiments and interpreted them with their own biases. Detractors of the theory of spontaneous generation were just as guilty of bias but in the opposite direction. It is somewhat surprising that Pasteur and Spallanzoni did not get growth in their cultures since the sterilization conditions they used would often not kill endospores . Luck certainly played a role. It is important to keep in mind that the discipline of science is performed by humans with all the fallibility and bias inherent in the species. Only the self-correcting nature of the practice reduces the impact of these biases on generally held theories. Spontaneous generation was a severe test of scientific experimentation because it was such a seductive and widely held belief. Yet, even spontaneous generation was overthrown when the weight of careful experimentation argued against it. Table 1.3 lists important events in the spontaneous generation debate.

Table 1.3 Events in spontaneous generation

Key takeaways.

• For many centuries many people believed in the concept of spontaneous generation, the creation of life from organic matter.
• Francesco Redi disproved spontaneous generation for large organisms by showing that maggots arose from meat only when flies laid eggs in the meat.
• Spontaneous generation for small organisms again gained favor when John Needham showed that if a broth was boiled (presumed to kill all life) and then allowed to sit in the open air, it became cloudy.
• Louis Pasteur ended the debate with his famous swan-neck flask experiment, which allowed air to contact the broth. Microbes present in the dust were not able to navigate the tortuous bends in the neck of the flask.

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Francesco Redi: Founder of Experimental Biology

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Francesco Redi was an Italian naturalist, physician, and poet. Besides Galileo, he was one of the most important scientists who challenged Aristotle 's traditional study of science. Redi gained fame for his controlled experiments. One set of experiments refuted the popular notion of spontaneous generation—a belief that living organisms could arise from nonliving matter. Redi has been called the "father of modern parasitology" and the "founder of experimental biology".

Birth : February 18, 1626, in Arezzo, Italy

Death : March 1, 1697, in Pisa Italy, buried in Arezzo

Nationality : Italian (Tuscan)

Education : University of Pisa in Italy

Published Work s: Francesco Redi on Vipers ( Osservazioni intorno alle vipere) , Experiments on the Generation of Insects ( Esperienze Intorno ​alla Generazione degli Insetti) , Bacchus in Tuscany ( Bacco in Toscana )

Major Scientific Contributions

Redi studied  venomous snakes to dispel popular myths about them. He demonstrated that it is not true that vipers drink wine, that swallowing snake venom is toxic, or that venom is made in a snake's gallbladder. He found that venom was not poisonous unless it entered the bloodstream and that the progression of venom in the patient could be slowed if a ligature was applied. His work paved the foundation for the science of toxicology .

Flies and Spontaneous Generation

One of Redi's most famous experiments investigated spontaneous generation . At the time, scientists believed in the Aristotelian idea of abiogenesis , in which living organisms arose from non-living matter. People believed rotting meat spontaneously produced maggots over time. However, Redi read a book by William Harvey on generation in which Harvey speculated that insects, worms, and frogs might arise from eggs or seeds too tiny to be seen. Redi devised and performed the now-famous experiment in which six jars, half left in open air and half covered with fine gauze that permitted air circulation but kept out flies, were filled with either an unknown object, a dead fish, or raw veal. The fish and veal rotted in both groups, but maggots only formed in the jars open to air. No maggots developed in the jar with the unknown object.

He performed other experiments with maggots, including one where he placed dead flies or maggots in sealed jars with meat and observed living maggots did not appear. However, when he placed living flies were placed in a jar with meat, maggots did appear. Redi concluded maggots came from living flies, not from rotting meat or from dead flies or maggots.

The experiments with maggots and flies were important not only because they refuted spontaneous generation, but also because they used control groups , applying the scientific method to test a hypothesis.

Parasitology

Redi described and drew illustrations of over one hundred parasites, including ticks, nasal flies, and the sheep liver fluke. He drew a distinction between the earthworm and the roundworm , which were both considered to be helminths prior to his study. Francesco Redi performed chemotherapy experiments in parasitology, which were noteworthy because he used an experimental control. In 1837, Italian zoologist Filippo de Filippi named the larval stage of the parasitic fluke "redia" in honor of Redi.

Redi's poem "Bacchus in Tuscany" was published after his death. It is considered among the best literary works of the 17th century. Redi taught the Tuscan language, supported the writing of a Tuscan dictionary, was a member of literary societies, and published other works.

Redi was a contemporary of Galileo, who faced opposition from the Church. Although Redi's experiments ran contrary to the beliefs of the time, he did not have the same sort of problems. This may well have been because of the different personalities of the two scientists. While both were outspoken, Redi did not contradict the Church. For example, in reference to his work on spontaneous generation, Redi concluded  omne vivum ex vivo  ("All life comes from life").

It's interesting to note that despite his experiments, Redi believed spontaneous generation could occur, for instance, with intestinal worms and gall flies.

Altieri Biagi; Maria Luisa (1968). Lingua e cultura di Francesco Redi, medico . Florence: L. S. Olschki.

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