louis pasteur swan neck experiment explained

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Pasteur's Experiment

The steps of Pasteur's experiment are outlined below:

First, Pasteur prepared a nutrient broth similar to the broth one would use in soup.

Next, he placed equal amounts of the broth into two long-necked flasks. He left one flask with a straight neck. The other he bent to form an "S" shape.

Then he boiled the broth in each flask to kill any living matter in the liquid. The sterile broths were then left to sit, at room temperature and exposed to the air, in their open-mouthed flasks.

After several weeks, Pasteur observed that the broth in the straight-neck flask was discolored and cloudy, while the broth in the curved-neck flask had not changed.

He concluded that germs in the air were able to fall unobstructed down the straight-necked flask and contaminate the broth. The other flask, however, trapped germs in its curved neck,­ preventing them from reaching the broth, which never changed color or became cloudy.

If spontaneous generation had been a real phenomenon, Pasteur argued, the broth in the curved-neck flask would have eventually become reinfected because the germs would have spontaneously generated. But the curved-neck flask never became infected, indicating that the germs could only come from other germs.

Pasteur's experiment has all of the hallmarks of modern scientific inquiry. It begins with a hypothesis and it tests that hypothesis using a carefully controlled experiment. This same process — based on the same logical sequence of steps — has been employed by scientists for nearly 150 years. Over time, these steps have evolved into an idealized methodology that we now know as the scientific method. After several weeks, Pasteur observed that the broth in the straight-neck flask was discolored and cloudy, while the broth in the curved-neck flask had not changed.

Let's look more closely at these steps.

Please copy/paste the following text to properly cite this HowStuffWorks.com article:

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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3. The Cell

3.1 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

CLINICAL FOCUS: Part 1

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 (“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 moulded, 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 2 ).

• 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?

Key Takeaways

• The theory of spontaneous generation states that life arose from nonliving matter. It was a long-held belief dating back to Aristotle and the ancient Greeks.
• Experimentation by Francesco Redi in the 17th century presented the first significant evidence refuting spontaneous generation by showing that flies must have access to meat for maggots to develop on the meat. Prominent scientists designed experiments and argued both in support of (John Needham) and against (Lazzaro Spallanzani) spontaneous generation.
• Louis Pasteur is credited with conclusively disproving the theory of spontaneous generation with his famous swan-neck flask experiment. He subsequently proposed that “life only comes from life.”

Multiple Choice

Fill in the blank, short answer.

• Explain in your own words Pasteur’s swan-neck flask experiment.
• Explain why the experiments of Needham and Spallanzani yielded in different results even though they used similar methodologies.

Critical Thinking

• What would the results of Pasteur’s swan-neck flask experiment have looked like if they supported the theory of spontaneous generation?

Media Attributions

• OSC_Microbio_03_01_Rediexpt
• https://link.springer.com/content/pdf/10.1007%2Fs10739-017-9494-7.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. ↵

Microbiology: Canadian Edition Copyright © 2019 by Wendy Keenleyside is licensed under a Creative Commons Attribution 4.0 International License , except where otherwise noted.

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The middle years 1862-1877

Louis Pasteur’s work raised a new set of research questions, such as " Where do fermentation agents come from ? " and " Do they originate from germs similar to themselves or do they appear spontaneously as explained by the spontaneous generation theory ? "

Spontaneous generation - the big debate

At the time the spontaneous generation theory was widely accepted in scientific circles. Louis Pasteur decided to approach the issue via his experimental method.

This required the use of swan-necked flasks. Water in the flask was brought to the boil for a few minutes until the steam escaped from the open end of the flask. It was then left to cool. While cooling, the air entering the flask deposited dust and germs on the first bend. Although in contact with outside air the liquid remained unaltered because germs could not get through.

Louis Pasteur showed that microbes were omnipresent - in water, in air, on objects, on the skin – and that some were responsible for diseases.

After some memorable struggles against his opponents, notably the famous biologist and fierce defender of the spontaneous generation theory, Félix Pouchet, in his 1862 paper Louis Pasteur was able to claim that :

• airborne dust contained microorganisms which develop and multiply.
• even the most putrescible liquids remained unadulterated if kept away from air (and hence these microorganisms) after heating.

He recommended ways of preventing and fighting these germs, and thus the habits essential for  personal and social hygiene . This notably included the use of  aseptic procedure s, i.e. the various measures to be taken to prevent invasion of live tissue or inert environments by exogenous microorganisms or viruses. He advocated the importance of  sterilization of linen and dressings, passing instruments through a flame and clean hands . These recommendations led to the widespread advent of modern surgery.

So how does fermentation work ?

But Louis Pasteur still had ferments in mind. He pondered on fermentation and how ferments work. While studying butyric fermentation he discovered a new class of living organisms capable of living without air.

He used the term " anaerobic " to describe ferments able to live without air and " aerobic " for microorganisms requiring the presence of free oxygen to grow.

He came to the conclusion that fermentation is the consequence of life without air.

He applied his microbiological method to industry and agriculture to eradicate ancient diseases affecting crops and products.

  To the rescue of industry and agriculture

He studied the formation of vinegar and the conversion of alcohol into acetic acid by Mycoderma aceti, which fixes oxygen from the air onto the alcohol. He showed vinegar makers how to produce vinegar of consistent quality by avoiding contamination by harmful mycoderma.  

Wine diseases

Wine was France flagship industry and a difficult business in many respects. Winemakers had difficulty guaranteeing the quality of their production which was affected by diseases of no known cause or cure. The crisis was nothing new but risked damaging exports and above all trade agreements in place with England. Emperor Napoleon III called on Louis Pasteur to seek a solution.   First he showed that each wine disease was due to a particular ferment.   He developed a protocol to fight the diseases, heating the wine to between 55°C and 60°C, a temperature at which it does not deteriorate and its bouquet is preserved. This method is now known worldwide as pasteurization .  

Just like wine, beer is infected by microorganisms transmitted by airborne dust. Louis Pasteur taught brewers to preserve the wort from the impurities and to heat the beer to 55° to prevent disease.

Silkworm diseases

In 1865, disease hit the silk industry. In France, this posed a threat to the economy of an entire region and the disease spread further afield to other silk-producing countries such as Italy, Austria and Asia Minor.   Louis Pasteur discovered that silkworms were affected by two diseases - silkworm nosema disease and flacherie.

Under the microscope, Louis Pasteur noticed that the worms with nosema disease developed shiny corpuscles, and showed that the disease was both hereditary and contagious.   He developed the cellular egg production method to enable the preservation of healthy silkworm eggs. He isolated the female moths to allow them to lay their eggs separately. After laying, he ground the female moths and examined them under the microscope. If the shiny corpuscles were observed he destroyed the eggs, otherwise he kept them for breeding.   As for flacherie, he introduced the "specific terrain" concept, i.e. the physiological condition of the infected host favoring outbreak of the disease. A few hygiene rules, good ventilation and quarantine of the suspect batches sufficed to prevent contamination.   These simple processes saved the silk industry from doom. But the research was of considerable value, paving the way for the study of contagious diseases. For the first time problems of heredity and contagion were scientifically proven and prophylaxis rules were established.   The time had now come for Louis Pasteur to address human diseases.

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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 Joh n Tyndall showed 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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Fermentation and putrefaction were often perceived as being spontaneous phenomena, a perception stemming from the ancient belief that life could generate spontaneously. During the 18th century the debate was pursued by the English naturalist and Roman Catholic divine John Turberville Needham and the French naturalist Georges-Louis Leclerc, count de Buffon . While both supported the idea of spontaneous generation , Italian abbot and physiologist Lazzaro Spallanzani maintained that life could never spontaneously generate from dead matter. In 1859, the year English naturalist Charles Darwin published his On the Origin of Species , Pasteur decided to settle this dispute. He was convinced that his germ theory could not be firmly substantiated as long as belief in spontaneous generation persisted. Pasteur attacked the problem by using a simple experimental procedure. He showed that beef broth could be sterilized by boiling it in a “swan-neck” flask, which has a long bending neck that traps dust particles and other contaminants before they reach the body of the flask. However, if the broth was boiled and the neck of the flask was broken off following boiling, the broth, being reexposed to air, eventually became cloudy, indicating microbial contamination. These experiments proved that there was no spontaneous generation, since the boiled broth, if never reexposed to air, remained sterile . This not only settled the philosophical problem of the origin of life at the time but also placed on solid ground the new science of bacteriology , which relied on proven techniques of sterilization and aseptic manipulation.

In 1862 Pasteur was elected to the Académie des Sciences , and the following year he was appointed professor of geology, physics , and chemistry at the École des Beaux-Arts (School of Fine Arts). Shortly after this, Pasteur turned his attention to France’s silkworm crisis. In the middle of the 19th century, a mysterious disease had attacked French silkworm nurseries. Silkworm eggs could no longer be produced in France, and they could not be imported from other countries, since the disease had spread all over Europe and had invaded the Caucasus region of Eurasia, as well as China and Japan. By 1865 the silkworm industry was almost completely ruined in France and, to a lesser extent, in the rest of western Europe. Pasteur knew virtually nothing about silkworms, but, upon the request of his former mentor Dumas, Pasteur took charge of the problem, accepting the challenge and seizing the opportunity to learn more about infectious diseases . He soon became an expert silkworm breeder and identified the organisms that caused the silkworm disease. After five years of research, he succeeded in saving the silk industry through a method that enabled the preservation of healthy silkworm eggs and prevented their contamination by the disease-causing organisms. Within a couple of years, this method was recognized throughout Europe; it is still used today in silk-producing countries.

In 1867 Pasteur resigned from his administrative duties at the École Normale Supérieure and was appointed professor of chemistry at the Sorbonne, a university in Paris . Although he was partially paralyzed (left hemiplegia ) in 1868, he continued his research. For Pasteur, the study of silkworms constituted an initiation into the problem of infectious diseases, and it was then that he first became aware of the complexities of infectious processes. Accustomed as he was to the constancy and accuracy of laboratory procedures, he was puzzled by the variability of animal life, which he had come to recognize through his observation that individual silkworms differed in their response to disease depending on physiological and environmental factors. By investigating these problems, Pasteur developed certain practices of epidemiology that served him well a few years later when he dealt with animal and human diseases.

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3 1.2 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

CLINICAL FOCUS: Part 1

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 (“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 moulded, 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 1.10 ). 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 1.11 ).

• 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 1.12 ). 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?

Key Takeaways

• The theory of spontaneous generation states that life arose from nonliving matter. It was a long-held belief dating back to Aristotle and the ancient Greeks.
• Experimentation by Francesco Redi in the 17th century presented the first significant evidence refuting spontaneous generation by showing that flies must have access to meat for maggots to develop on the meat. Prominent scientists designed experiments and argued both in support of (John Needham) and against (Lazzaro Spallanzani) spontaneous generation.
• Louis Pasteur is credited with conclusively disproving the theory of spontaneous generation with his famous swan-neck flask experiment. He subsequently proposed that “life only comes from life.”

Multiple Choice

Fill in the blank, short answer.

• Explain in your own words Pasteur’s swan-neck flask experiment.
• Explain why the experiments of Needham and Spallanzani yielded in different results even though they used similar methodologies.

Critical Thinking

• What would the results of Pasteur’s swan-neck flask experiment have looked like if they supported the theory of spontaneous generation?
• https://link.springer.com/content/pdf/10.1007%2Fs10739-017-9494-7.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. ↵

DeSales Microbiology Copyright © 2022 by DeSales University & Dr. Dia Beachboard is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License , except where otherwise noted.

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