The Science of Tomato Flavors

Don’t judge a tomato by its appearance. You may salivate when thinking about the luscious large red tomatoes you just purchased in your grocery store, only to find out that they are extremely bland and lack flavor once you actually bite into them after preparing the salad you had been looking forward to all day. You are not alone. Many consumers complain about the growing blandness of fruits. Up until a few decades ago, it was rather challenging to understand the scientific basis of fruit flavors. Recent biochemical and molecular studies of fruits now provide a window into fruit flavors and allow us to understand the rise of blandness.

In a recent article, the scientists Harry Klee and Denise Tieman at the University of Florida summarize some of the most important recent research on the molecular biology of fruit flavors, with a special emphasis on tomatoes. Our perception of “flavor” primarily relies on two senses – taste and smell. Taste is perceived by taste receptors in our mouth, primarily located on the tongue and discriminates between sweet, sour, salty, bitter and savory. The sensation of smell (also referred to as “olfaction”), on the other hand, has a much broader catalog of perceptions. There are at least 400 different olfactory receptors present in the olfactory epithelium – the cells in the nasal passages which perceive smells – and the combined activation of various receptors can allow humans to distinguish up to 1 trillion smells. These receptors are activated by so-called volatile organic compounds or volatiles, a term which refers to organic molecules that are vaporize in the mouth when we are chewing the food and enter our nasal passages to activate the olfactory epithelium. The tremendous diversity of the olfactory receptors thus allows us to perceive a wide range of flavors. Anybody who eats food while having a cold and a stuffy nose will notice how bland food has become, even though the taste receptors on the tongue remain fully functional.

When it comes to tomato flavors, research has shown that consumers clearly prefer “sweetness”. One obvious determinant of sweetness is the presence of sugars such as glucose or fructose in tomatoes which are sensed by the taste receptors in the mouth. But it turns out that several volatiles are critical for the perception of “sweetness” even though they are not sugars but instead activate the smell receptors in the olfactory epithelium. 6-Methyl-5-hepten-2-one, 1-Nitro-2-phenylethane, Benzaldehyde and 2-Phenylethanol are examples of volatiles that enhance the positive flavor perceived by consumers, whereas volatiles such as Eugenol and Isobutyl acetate are perceived to contribute negatively towards flavor. Interestingly, the same volatiles can have no effect or even the opposite effect on flavor perception when present in other fruits. Therefore, it appears that for each fruit, the sweetness flavor is created by the basic taste receptors which sense sugar levels as well as a symphony of smell sensations activated by a unique pattern of volatiles. But just like instruments play defined yet interacting roles in an orchestra, the effect of volatiles on flavor depends on the presence of other volatiles.

This complexity of flavor perception explains why it is so difficult to define flavor. The story becomes even more complicated because individuals have different thresholds for olfactory receptor activation. Furthermore, even the volatiles linked with a positive flavor perception – either by enhancing flavor intensity or letting the consumer sense a greater “sweetness” then actually present based on sugar levels – may have varying effects when they reach higher levels. Thus, it is very difficult to breed the ideal tomato that will satisfy all consumers. But why is there this growing sense that fruits such as tomatoes are becoming blander? Have we simply not tried enough tomato cultivars? A cultivar is a plant variety that has been bred over time to create specific characteristics, and one could surmise that with hundreds or even thousands of tomato cultivars available, each of us might identify a distinct cultivar that we find most flavorful. The volatiles are generated by metabolic enzymes encoded by genes and differences between the flavor of distinct cultivars is likely a reflection of differences in gene expression for the enzymes that regulate sugar metabolism or volatiles generation.

The problem, according to Klee and Tieman, is that the customers of tomato breeders are tomato growers and not the consumers who garnish their salads or create tomato-based masalas. The goal of growers is to maximize shelf-life, appearance, disease-resistance, yield and uniformity. Breeders focus on genetically manipulating tomato strains to maximize these characteristics. The expression GMO (genetically modified organism) describes the use of modern genetic technology to modify individual genes in crops and often provokes a litany of attacks and criticisms by anti-GMO activists who fear potential risks of such genetic interventions. However, the genetic breeding and manipulation of cultivars has been occurring for centuries or even millennia using traditional low tech methods but these do not seem to provoke much criticism by anti-GMO activists. Even though there is a theoretical risk that modern genetic engineering tools could pose a health risk, there is no scientific evidence that this is actually the case. Instead, one could argue that targeted genetic intervention may be more precise using modern technologies than the low-tech genetic breeding manipulations that have led to the creation of numerous cultivars, many of whom carry the “organic, non-GMO” label.

Klee and Tieman argue that consumers prefer flavor, variety and nutrition instead of the traditional goals of growers. The genetic and biochemical analysis of tomato cultivars now offers us a unique insight into the molecular components of flavor and nutrition. Scientists can now analyze each cultivar that has been generated over the past centuries using the low-tech genetic manipulation of selective breeding and inform consumers as to their flavor footprint. Alternatively, one could also use modern genetic tools such as genome editing and specifically modify flavor components while maintaining disease-resistance and high nutritional value of crops such as tomatoes. The key to making informed, rational decisions is to provide consumers comprehensive information based on scientific evidence as to the nutritional value and flavor of fruits, as well as the actual risks of genetically modifying crops using traditional low tech methods such as selective breeding and grafting or newer methods which involve genome editing.


Klee, H. J & Denise M. Tieman (2018). The genetics of fruit flavour preferencesNature Reviews Genetics, (published online March 2018)

Note: An earlier version of this article was first published on the 3Quarksdaily blog.


Blissful Ignorance: How Environmental Activists Shut Down Molecular Biology Labs in High Schools

Hearing about the HannoverGEN project made me feel envious and excited. Envious, because I wish my high school had offered the kind of hands-on molecular biology training provided to high school students in Hannover, the capital of the German state of Niedersachsen. Excited, because it reminded me of the joy I felt when I first isolated DNA and ran gels after restriction enzyme digests during my first year of university in Munich. I knew that many of the students at the HannoverGEN high schools would be similarly thrilled by their laboratory experience and perhaps even pursue careers as biologists or biochemists.

What did HannoverGEN entail? It was an optional pilot program initiated and funded by the state government of Niedersachsen at four high schools in the Hannover area. Students enrolled in the HannoverGEN classes would learn to use molecular biology tools typically reserved for college-level or graduate school courses in order to study plant genetics. Some of the basic experiments involved isolating DNA from cabbage or how learning how bacteria transfer genes to plants, more advanced experiments enabled the students to analyze whether or not the genome of a provided maize sample had been genetically modified. Each experimental unit was accompanied by relevant theoretical instruction on the molecular mechanisms of gene expression and biotechnology as well as ethical discussions regarding the benefits and risks of generating genetically modified organisms (“GMOs”). The details of the HannoverGEN program are only accessible through the the Wayback Machine Internet archive because the award-winning educational program and the associated website were shut down in 2013 at the behest of German anti-GMO activist groups, environmental activists, Greenpeace, the Niedersachsen Green Party and the German organic food industry.

Why did these activists and organic food industry lobbyists oppose a government-funded educational program which improved the molecular biology knowledge and expertise of high school students? A press release entitled “Keine Akzeptanzbeschaffung für Agro-Gentechnik an Schulen!” (“No Acceptance for Agricultural Gene Technology at Schools“) in 2012 by an alliance representing “organic” or “natural food” farmers accompanied by the publication of a critical “study” with the same title (PDF), which was funded by this alliance as well as its anti-GMO partners, gives us some clues. They feared that the high school students might become too accepting of biotechnology in agriculture and that the curriculum did not sufficiently highlight all the potential dangers of GMOs. By allowing the ethical discussions to not only discuss the risks but also mention the benefits of genetically modifying crops, students might walk away with the idea that GMOs could be beneficial for humankind. The group believed that taxpayer money should not be used to foster special interests such as those of the agricultural industry which may want to use GMOs.

A response by the University of Hannover (PDF), which had helped develop the curriculum and coordinated the classes for the high school students, carefully analyzed the complaints of the anti-GMO activists. The author of the anti-HannoverGEN “study” had not visited the HannoverGEN laboratories, nor had he had interviewed the biology teachers or students enrolled in the classes. In fact, his critique was based on weblinks that were not even used in the curriculum by the HannoverGEN teachers or students. His analysis ignored the balanced presentation of biotechnology that formed the basis of the HannoverGEN curriculum and that discussing potential risks of genetic modification was a core topic in all the classes.

Unfortunately, this shoddily prepared “study” had a significant impact, in part because it was widely promoted by partner organizations. Its release in the autumn of 2012 came at an opportune time for political activists because Niedersachsen was about to have an election. Campaigning against GMOs seemed like a perfect cause for the Green Party and a high school program which taught the use of biotechnology to high school students became a convenient lightning rod. When the Social Democrats and the Green Party formed a coalition after winning the election in early 2013, nixing the HannoverGEN high school program was formally included in the so-called coalition contract. This is a document in which coalition partners outline the key goals for the upcoming four year period. When one considers how many major issues and problems the government of a large German state has to face, such as healthcare, education, unemployment or immigration, it is mind-boggling that de-funding a program involving only four high schools received so much attention that it needed to be anchored in the coalition contract. In fact, it is a testimony to the influence and zeal of the anti-GMO lobby.

Once the cancellation of HannoverGEN was announced, the Hannover branch of Greenpeace also took credit for campaigning against this high school program and celebrated its victory. The Greenpeace anti-GMO activist David Petersen said that the program was too cost intensive because equipping high school laboratories with state-of-the-art molecular biology equipment had already cost more than 1 million Euros. The previous center-right government which had initiated the HannoverGEN project was planning on expanding the program to even more high schools because of the program’s success and national recognition for innovative teaching. According to Petersen, this would have wasted even more taxpayer money without adequately conveying the dangers of using GMOs in agriculture.

The scientific community was shaken up by the decision of the new Social Democrat-Green Party coalition government in Niedersachsen. This was an attack on the academic freedom of schools under the guise of accusing them of promoting special interests while ignoring that the anti-GMO activists were representing their own special interests. The “study” attacking HannoverGEN was funded by the lucrative “organic” or “natural food” food industry! Scientists and science writers such as Martin Ballaschk or Lars Fischer wrote excellent critical articles stating that squashing high-quality, hand-on science programs could not lead to better decision-making. How could ignorant students have a better grasp of GMO risks and benefits than those who receive relevant formal science education and thus make truly informed decisions? Sadly, this outcry by scientists and science writers did not make much of a difference. It did not seem that the media felt this was much of a cause to fight for. I wonder if the media response would have been just as lackluster if the government had de-funded a hands-on science lab to study the effects of climate change.

In 2014, the government of Niedersachsen then announced that they would resurrect an advanced biology laboratory program for high schools with the generic and vague title “Life Science Lab”. By removing the word “Gen” from its title which seems to trigger visceral antipathy among anti-GMO activists, de-emphasizing genome science and by also removing any discussion of GMOs from the curriculum, this new program would leave students in the dark about GMOs. Ignorance is bliss from an anti-GMO activist perspective because the void of scientific ignorance can be filled with fear.

From the very first day that I could vote in Germany during the federal election of 1990, I always viewed the Green Party as a party that represented my generation. A party of progressive ideas, concerned about our environment and social causes. However, the HannoverGEN incident is just one example of how the Green Party is caving in to ideologies, thus losing its open-mindedness and progressive nature. In the United States, the anti-science movement, which attacks teaching climate change science or evolutionary biology at schools, tends to be rooted in the right wing political spectrum. Right wingers or libertarians are the ones who always complain about taxpayer dollars being wasted and used to promote agendas in schools and universities. But we should not forget that there is also a different anti-science movement rooted in the leftist and pro-environmental political spectrum – not just in Germany. As a scientist, I feel that it is becoming increasingly difficult to support the Green Party because of its anti-science stance.

I worry about all anti-science movements, especially those which attack science education. There is nothing wrong with questioning special interests and ensuring that school and university science curricula are truly balanced. But the balance needs to be rooted in scientific principles, not political ideologies. Science education has a natural bias – it is biased towards knowledge that is backed up by scientific evidence. We can hypothetically discuss dangers of GMOs but the science behind the dangers of GMO crops is very questionable. Just like environmental activists and leftists agree with us scientists that we do not need to give climate change deniers and creationists “balanced” treatment in our science curricula, they should also accept that much of the “anti-GMO science” is currently more based on ideology than on actual scientific data. Our job is to provide excellent science education so that our students can critically analyze and understand scientific research, independent of whether or not it supports our personal ideologies.


Note: An earlier version of this article was first published on the 3Quarksdaily blog.